Breeding Techniques and Selection for Breeding of the Honeybee-Online-V1

BREEDING TECHNIQUES AND SELECTION FOR BREEDING OF THE HONEYBEE FRIEDRICH RUTTNER

Breeding Techniques and Selection for Breeding of the Honeybee

An Introduction to the Rearing of Queens, the Conduct of Selection Procedures and the Operation of Mating Stations

Translated by Ashleigh and Eric Milner

Published by The British Isles Bee Breeders Association by arrangement with Ehrenwirth Verlag, Munich British Library Cataloguing in Publication Data Ruttner, Friedrich Breeding techniques and selection for breeding of the honeybee. 1. Western Europe. Livestock: Honey bees. Breeding I. Title II. British Isles Bee Breeders' Association III. Zuchttechnik und Zuchtauslese bei der Beine. English 638'. 1

ISBN 0-905369-07-6

This edition Published by The British Isles Bee Breeders Association by arrangement with Ehrenwirth Verlag, Munich

Translated by Ashleigh and Eric Milner

Printed in England by G Beard & Son Ltd, Brighton Contents Translators' Foreword 4 Preface to the 6th Edition 7 Introduction 8

I Rearing of Queens and Drones 11 1. Rearing of Queens 12 2. Rearing of Drones 41 3. Co-ordination of Drone and Queen-rearing 44

II Selection for performance 45 1. What is Breeding? 45 2. The Breeding of Honeybees 46 3. Results of Selective Breeding 48 4. The Evaluation of Performance 53 5. The Honeybee Family 59 6. Breeding Programmes 63

III Control of Mating 79 1. The Mating Apiary 79 2. The Pure-Breeding Area 80 3. The Mating Station 81

IV The Races and Strains of Honeybees of Central Europe 96 1. Distribution 96 2. Comparison of Carnica and Mellifera 98 3 The Incompatibility of the Honeybee Races 100 4. Cross Breeding 100 5. How do Carnica and Mellifera differ from each other? 103 6. Breeding Strains 104 7. Other Races of Significance for Breeding 108

V The Assessment of the Physical Characters 111 1. Sampling 111 2. Instruments 112 3. The Racial Characters 112 4. Interpretation of the Character-data 137 5. The Physical Characters of the Races of the Honeybee 138 6. Measurement of the Proboscis 141

Notes 146

INDEX 149 Translators' Foreword

Good wine needs no bush, and a book by Dr. Ruttner needs no commenda- tion. His name is known and respected among scientific beekeepers in five continents. He has supplied us with the following information and advice: "From the time when Europe's beekeepers detected the existence of various races of honeybees, importation of foreign stock and the inevitable hybridisation of different geographic races began, with all the well known consequences for the behaviour of the colonies. Early attempts were made to re-establish a more manageable bee, but selection programmes and planned matings have produced substantial results only since the characteristics of races, and mating biology were better known and simple biometrical methods, applicable by beekeepers were also available. Since 1948 the author has devoted his efforts towards this goal, concen- trating mainly on the discrimination of the two dark European races: the Black Bee of Central and Western Europe, and Carniolans. The frequent hybrids between these races can be recognised only by special methods, which are now adopted by beekeepers all over Germany and Austria, and also in other countries. They help to keep the two races accurately separated, even within the same country, e.g. in Norway. Hundreds of week-end classes were - and are - held to make the beekeepers familiar with these methods. The procedures described in this booklet were first introduced while the author was at the Breeding Station, Lunz am See, Austria, and were completed later at the Apicultural Institute, Oberursel, West Germany. An elaborate selection and testing programme, effective queen rearing and mating control, with a check up of the result, obtained by morphomet- ries, constitute a unit. Forty years after the start, clear progress obtained with this breeding programme using Carniolan strains, is clearly demonstrated. At the same time the almost vanished native Black Bee of the Tyrolean Alps was preserved and improved with the same methods". We hope that this book will encourage the breeding and improvement of our native Dark Bee in these islands. Although in these pages it is compared unfavourably with the Carniolan, we must remember that the Carniolan has had the benefit of selection and improvement for nearly a century, further advanced by more than forty years of scientific work while the Dark Bee has been largely neglected. We hope that, profiting by the skill and knowledge conveyed in these pages, our readers will remedy this defect. We are further encouraged by Dr. Ruttner's statement that while the Carniolan Bee is the bee for the continental climate, the Dark Bee is the bee for the Atlantic seaboard with its uncertain weather and in particular its erratic springs. But how can we improve or breed from an extinct race? Since the 1920s it has been constantly asserted that the British Bee was exterminated by the Isle of Wight Disease. Is this a fact? Is it even probable? Recently, an attempt was made to rid the country of rabbits by the introduction of Myxomatosis. Great numbers died, but some were resistant, even when

4 new strains of the disease were introduced, and as is well known they have multiplied again. It seems highly probable that something similar may have happened to our native bees. The bees of today are descendants of the immune or resistant survivors. For whatever reason, many have certainly survived. We have received samples from Scotland, Wales and different parts of the north and west of England which conform closely to the morphometric standards of Apis mellifera mellifera, and are reported to show the traditional characters of the British bee: compact brood nest, always with reserves of honey and pollen; early breeding and robust wintering. This has been gratifying but not surprising. The native bee has become aclimatised since the last Ice Age. It winters well (A Scottish beekeeper said in 1986 : Bad winter? those who keep these bees don't know there's been a bad winter. Those who import bees daren't face the bank manager). Native bees rear drones and queens earlier in the year than imported bees do, and they can fly at a lower temperature than the foreign bees can tolerate. So with early mating they enjoy ecological isolation and can maintain racial purity. Anxiety is sometimes expressed that the native bee will not develop in Spring in time for the early nectar flow. From our experience (in Notts, and Yorks) we can state that if our colonies have been well prepared for winter, with adequate stores, they will usually build up to great strength in time to bring in a good harvest from the Rape in April and be ready for swarming or queen rearing early in May. As with all bees, there is variation, and a programme of selection can improve performance in this respect. We have also observed that some queens do not develop their full potential till the second year. The need for improvement is urgent. In our thickly populated country it is of prime importance that our bees should be quiet and gentle. Experience confirms Dr. Ruttner's teaching that only bees of pure race can be relied on to transmit desirable qualities to their offspring. To achieve this for the country as a whole breeders will need the conditions described in this book; isolated mating stations, pure race areas or instrumental insemination. Till these conditions are attained we shall have to rely on the qualities which have enabled our bees to survive in a pure condition, good wintering, early queen rearing and low temperature flight. The techniques for the biometrical identification of bees and for selective breeding, lucidly set out by Dr. Ruttner have been successfully carried out by groups of workers in this country during the last six years and have been found to be within the competence of many beekeepers. Dr. Ruttner authorised us to make, in the translation, such adaptations as we thought necessary to the conditions of British Apiculture, but we thought it wiser and fairer to the reader to adhere as closely as we could to Dr. Ruttner's own words and to confine our remarks to the notes. Our heartiest thanks are offered to Dr. Ruttner for encouraging the publication of our translation, and for supplying the new material to bring it into line with his new edition.

5 We also wish to thank: Bernard Mobus, N.D.B., till recently C.B.I, for N.E. Scotland; Dr. J. van Praagh, of the Institute for Bee Research, Celle; Dr. V. Maul, of the Bee Breeding Institute, Kirchhain; Dr. F. Schaper, of the Institute for Bee Breeding, Erlangen; and Frau Kuhnert, of the Institute for Bee Research, Oberursel. The readers' thanks are due to John Dews, who asked for the translation of two pages, then for more and more till the whole was done and then urged us on to publication, also to Eric Foster for great care with proof reading, and to Albert Knight, Secretary of BIBBA, for painstaking attention to the diagrams and special thanks to Mrs. Susan Grice, without whose unfailing patience and skill with a word processor this work would never have come to fruition. Lastly we mention the debt of British Beekeeping to the late Beowulf Cooper, without whose work, and his part in the foundation of BIBBA we should never have become acquainted with German Beekeeping, or had the privilege of meeting Dr. Ruttner.

6 Preface to the Sixth Edition These instructions for the rearing of queens, for selection procedures, and for the management of Mating Stations, give a concise summary of the experience which my brother, Dipl. Ing. Hans Ruttner (ob. Lunz am See 1979), my col- league for many years, and myself, were able to assemble from many years in practical breeding work, during training courses on selection breeding and by the evaluation of a very large quantity of breeding material. Emphasis has been placed on practical application and on a clear presentation. Consequently I have included only those methods and breeding principles which I have per- sonally tested and proved over many years. Each new edition of this work which has become necessary since its first appearance 25 years ago, has been based on an extension of knowledge and experience. It bears witness to the successes which can result from a vigorous combination of research and practical application. The preservation and improvement of the Carnica race only became possible through the study of the racial physical-characters of this bee in its country of origin, together with the biology of its mating on the one hand and on the other hand the precise measurement of the external physical characters, and the productivity of colonies, together with the setting up of reliable Mating Stations by the beekeeping industry. The success of this common endeavour is evidenced in ample numerical data: in the last 25 years the honey harvest of the breeding lines has continuously increased. The technique of identifying the derivation of colonies (and thereby the likely productive capacity) by measurement of the physical-racial characters - formerly often derided - is now increasingly employed, even in the U.S.A., in view of the menace of advancing "Africanised Bees". This English edition has been prepared from the new German edition. Selection for breeding is acknowledged as one of the supporting pillars of modern successful beekeeping. Every method that makes the selection process better and more accurate must be welcome. Only a beekeeper who regularly weighs, measures and records will achieve worthwhile results in the long term. A selection procedure in one form or another is therefore indispensable. Especially at a time when Varroatosis is rolling over Europe is it important that breeding successes already achieved be preserved and still further developed. It has long been recognised that queen renewal has an essential place in the armoury of successful management in a Varroa infested district. It should not by any means be ruled out that some day deliberate selective breeding will be introduced for this very purpose. Bee Master W. Gotz, with his ripe experience of breeding methods, has been of very great assistance in evaluating the breeding procedures. The organisation of the very great mass of data on the physical characters on which these instructions are based, was only made possible by the selfless assistance of my late wife, and the conscientious co-operation for very many years of my assistant, Agnes Mohr, who also prepared the illustrations for this edition. My thanks to Herr Dr. K. Dreher of Meyen, for permission to use Figures 32, 33 and 34.

Lunz am See March 1987 Friedrich Ruttner.

7 Introduction

All breeding begins with the choice of worthwhile breeding stock. The "value" of the breeding animals will first be determined by their own performance and qualities. But only a part of the selection process is involved here, since the gene assembly which will be transmitted to descendants interests the breeder much more than the immediately apparent performance, (i.e. the economic value of the colony). A performance that will not be inherited is without significance for the breeder. The essence of breeding is to transmit a certified, above average performance of individual animals to the greatest possible number of descendants, undiminished and as far as possible enhanced. There must therefore be a very clear distinction between the economic value and the breeding value of a stock. A "Blender",(I.1.) the product of crossbreeding has usually a very high economic value, but very rarely any breeding value. In the early stages of breeding, choice was made by "feel" using the so-called "Breeder's Glance" acquired through long familiarity with a particular strain. But the more technique and knowledge in breeding developed the greater was the endeavour to introduce objective and precise evaluation so as to make judgements which were generally more applicable and also more comparable. An investigation of breeding value carried out with exact, precisely defined procedures is designated "Selection". Selection procedures for honeybees are carried out in accordance with the following principles;-

1. Evaluation of Performance and Economically Important Qualities In order to compare performance all the usable data concerning honey yields and behaviour of all the colonies in the apiary must be considered (Record Cards!). Colonies without records of performance (for example, those with young queens) cannot be selected. At the most a provisional estimate on the basis of physical characters - "Preselection" - is possible for such a colony.

2. Proof of Pedigree Of course from both the father's and mother's sides. Data for the father's side can only be obtained from controlled mating (instrumental insemination or a thoroughly reliable Mating Station). The Pedigree Certificate should also include information about the performance of ancestors (parents) and sister stocks. The more clearly the average performance of the entire family exceeds the general average, so much the greater is the likeli- hood that this will also be true of the descendants.

3. Determination of the External Physical Characters The strains of bees that are bred in Central Europe belong without exception to one of two races, the grey Carniolan bee, and the Dark bee, which

can be unerringly distinguished from each other by body-structure, hairiness and wing venation. In addition, crossing with the Italian bee has still to be reckoned with, easily recognisable by the yellow rings on the abdomen. Experience over many years has shown that lasting results can only be obtained from breeding within a pure race, certainly not from breeding with the repeatedly crossed "local bees". By close examination of the physical characters we can ascertain whether bees conform to the type of one or the other of these races, and whether the bees of the stock under consideration are so uniform that similar progeny can be expected. At the same time it is also possible to make certain in many cases, whether the queen at the Mating Station has actually mated with drones that had been taken there as breeder drones. We have no other means at our disposal of checking matings, and only by strict and constant monitor- ing of characters can we eliminate in good time the "breakdowns" which we are never able to avoid entirely at our Mating Stations. Any breeding whatever is only of significance if one knows precisely what is being bred the whole of the time. In the breeding of large animals and plants this principle is so obvious that there is no need to waste words on it. With the tiny bee in which changes cannot be so easily detected the problem is more difficult. If in every generation scrupulously accurate observations of performance, behaviour and physical characters are not maintained, one can very soon be on the wrong road: in a very few years, the bees in the apiary may have very little in common with the material with which such a hopeful start was made. All the time and labour has been uselessly frittered away. Our bee is the smallest domestic animal. The consequence of this is that for assessing the physical characters we must get equipment such as a lens and forceps, and that considerable experience is needed to recognise the characteristic differences and to assess them correctly. Assessment of characters has simply and solely the purpose of deciding on the hereditable value of a queen which has an already proven performance. If the worker bees and drones of this colony consistently show the features of a particular race, then the performance will in all probability be inheritable. If, however, signs of hybridisation are apparent, one is dealing with a "Blender" and it would be waste of effort to breed from it. On no account however should selection for breeding be made by character selection pure and simple. At a good Mating Station a very "beautiful" bee with a quite uniform dark colour or with a very high Cubital Index can quickly be bred. But since it is unpractical at the same time to select for high performance the economic value of such a line is very quickly lost. Therein lies the danger of a one sided character selection, such as breeding for body colour only. There are no body characters that are directly related to performance qualities. The often attempted breeding for long tongues has never once been economically successful. To be sure, bees with longer tongues were obtained but with greatly reduced productivity.

9 We regard the natural strains and races of bees as a harmoniously bal- anced system with particular characters and qualities. This system con- sists of an enormous number of independent single elements which are not linked together but only loosely associated. Every crossing with another race disorganises these associations and creates a multitude of new, mainly less productive types. The change in the body characters is the outward sign of this disorganisation.

Which Colonies Can Be Selected? A full selection such as is demanded for breeder colonies is not possible with a single colony. Stocks can only be selected:- 1. Which have a recorded performance. Therefore, at the earliest a colony can only be selected after a complete production year of its queen. (Only in exceptional circumstances should a queen without records of performance be used for breeding). 2. For which there is evidence of performance of sister and parent colonies. 3. Whose queens have been mated at a Mating Station.

Inherited performance will show up in its full capacity only in colonies built up from a satisfactory queen. Therefore, the rearing of queens must first be dealt with before we turn to the actual breeding procedures.

10 Chapter I Rearing of Queens and Drones The aim of Selection is to identify and direct the heritable performance capabilities of the bee colonies. But along with the heritable there are numerous non-heritable factors which are involved in making performance a reality. The conditions under which the queens are reared and the mating process are very important. Although we speak of breeding selection with proper emphasis on heritable performance, we must not for- get that the highest performance can only be obtained from queens which have had in addition to the best ancestry and mating, the best rearing conditions as well. Anyone who undertakes Selection for Performance must take great pains to operate a method of rearing queens and drones that is quite flawless. The relative performance of a colony depends to a great extent on its numerical strength which in turn results from the egg-laying ability of the queen and on the nurse bees' devotion to the task of rearing. Both are controlled by heritable as well as environmental influences. The queen's ability to lay depends on her own inherited disposition but is influenced by the inherited disposition of her workers, which prepare the broodnest, feed the queen well or badly, and either nurse, or neglect, the eggs she lays. Of the external conditions, temperature and nectar-flow affect the workers most. At mating time the weather conditions and the number of flying drones in the area, determine the course of the mating and the amount of sperm transferred. Queens that are not well supplied with semen are bad layers, and are soon superseded. The important characters for reproduction and life span are developed by nourishment and temperature during the queen's larval and pupal stages. The technique of rearing must therefore ensure that optimal conditions prevail in this period for the formation of strong well developed queens. So it is clear that the technique of rearing must be dealt with in the framework of breeding selection. It can surely be assumed that a beekeeper who is taking an interest in Selective Breeding can already rear queens with confidence. So an account of elementary breeding precautions will be omitted in favour of an account of a few well tried methods, omitting all superfluous details. Above all, emphasis will be placed on those points where experience shows mistakes are most frequently made.

11 A detailed account of a number of well tested breeding methods together with their biological principles can be found in the Apimondia — Encyclopaedia: Queen Rearing: Biological Basis and Technical Information. (Obtainable from I.B.R.A.)

In breeding work the following principles apply;—

1. Efficiency; economy of time and material (i.e. of bees)

2. Attainment of the highest quality of breeding stock.

1. Rearing of Queens Very many breeding methods have been extolled but only a few have proved their worth in practice. We shall assess, according to the two principles stated above, which of the proposed methods are most likely to be successful. A breeding method must produce good queens, be simple and economical. The beekeeper can neither produce queens nor rear them himself; his contribution is restricted to offering larvae from a breeding colony to a rearing colony in such a way that they do the job for him in the best and most productive manner. The key position in rearing is therefore held by the nursing colony and its condition at the time. In my experience the influence of the nursing stock on the development of queens is of quite decisive significance and more important than most of the other matters we have to attend to in the rearing of queens. Unfortunately, we are only able to understand and influence a part of the factors which depend on the nursing colony.

The Ancestry of the Nurse Bees If we have been selecting breeding lines from a pure race for reluctance to swarm, we should not expect these colonies to be particularly suitable for nursing. Many establishments therefore regularly bring in stocks of "Heather Bees" (1.1.) for this purpose. This expedient has its drawbacks, on account of the continuous input of hybrid bees; furthermore, it is quite unnecessary. As a rule stocks headed by queens of the breeding line, but mated in the apiary, show very good nursing propensities because of the hybrid influence of the local bees.

The Condition of the Nursing Stock Vigorous colonies during the period of rapid growth are the most suitable for nursing, at any rate up to the time when swarm preparation begins. If the swarming mood is well advanced (open queen cells present) a good nursing mood is hardly likely to be created — biologically very under- standable. On the other hand immature stocks (those with an unsuitable bee : brood ratio) are ill adapted for nursing. I recommend as a minimum strength for a high quality nursing colony eight thickly covered combs literally dripping with bees. But if rearing is

12 to proceed, even under the worst weather conditions, then in practice the strongest possible colonies will always be used. Usually before the start of rearing the stock is given the opportunity to raise brood in two chambers to obtain the optimum number of nurses. One of the most effective expedients to prepare a stock for nursing is to crowd the bees into one chamber so that every corner is tightly packed, since the ratio of bees to space is of the utmost importance for nursing. It is particularly important to make a correct appraisal of the readiness of a colony for nursing at the start of the breeding season in Spring. At fruit-blossoming, the time of the early nectar flow, outstandingly good breeding conditions often occur on account of the weather and abundant supplies. In the desire for an early breeding batch, one of the strongest stocks is often dequeened — with resulting deep disappointment at the meagre low quality acceptances. What is the cause? The stock was not yet ripe for breeding. Particularly at this time of the year, ripeness for breeding does not depend solely on the number of bees; age structure plays a significant part. In Spring there are often still plenty of Winter bees present, old foragers which bring in the early nectar; but the number of young nurse bees is barely enough to meet the demands for larval food from the great number of larvae. The ratio, open brood : sealed brood, is an important pointer to a stock's "ripeness for rearing". Uniting two such "unripe" stocks does no good; at the best they can be turned into an artificial swarm and breeding started with it. But at this moment it is an expedient I would only recommend for the most compelling reason. Normally it is better to wait another 10 days. The bees inform us in another way that they are ready for queen rearing; by rearing drones. A colony without drones is not ripe for queen-rearing. The appearance of the first drones in Spring — which in the natural course of the bee's year is the earliest sign of swarming preparations, normally a short time before the appearance of the first queen cups — is for the breeder the signal for the start of queen rearing. The very earliest date, even before any drones have emerged, is when there is an area of sealed drone cells at least the size of the palm of one's hand. Even with strong nursing colonies we still emphasise the advice not to allow the rearing of more than 20-25 queen cells at the same time. The equation: double colony strength = double number of cells reared does not apply. This practical experience is only valid for Central European conditions. In other regions and with other races much larger batches can be raised without loss of quality

Known and Unknown Influences on Breeding Mood It is widely known that a light nectar flow has a marked effect on the work of a nursing colony. Complete drying up of the flow for a protracted period is naturally somewhat unfavourable, but in the earlier (expanding) part of the season it is not usually difficult by uninterrupted stimulative feeding to put the nursing colony in the right mood for breeding.

13 It is a different matter with a heavy nectar flow, especially a forest flow (honeydew). The bees come into such a flow fever that they neglect every- thing else, so that if the queen cells are given, only a few are accepted and these are badly tended. The result is few and badly developed queens. In such a situation it is better to move the target dates rather than carry out low grade breeding work. Should a forest flow be interrupted by bad weather, 1 or 2 days without a flow suffice to reawaken the breeding impulse. Acceptance and rearing during such a break in the flow is particularly good.

Bad and good nursing Colonies There are often failures for which we are not in a position to state the cause. Into a carefully prepared nursing colony, to all appearances just what it should be, the breeding material was introduced exactly according to rule. But the stock did not behave as it should; after 24 hours the edges of the cups were not drawn out and the larvae lay "naked" at the bottoms of the scarcely moistened cells. At the same time a neighbouring colony of the same strain had started to nurse a batch of grafts quite faultlessly. It is difficult to find a comprehensible explanation of these differences. If a colony does not want to nurse, we recommend that just one more "attempt at encouragement" be made; the stock is tested with "wild" queen cells, then given heavy stimulative feeding and even more strongly compressed than before. Then another batch is introduced. If this approach fails, it is better to prepare another nursing stock, rather than agonise over the first one, and perhaps in the end squeeze out of it nothing more than a few inferior queens. Reluctantly nursed and neglected larvae are much more harmful than rejected cells or a whole rejected batch. In the latter case the failure is immediately apparent and something must be done at once if queens are to be obtained. But if the larvae are still present, even if scarcely grown and very sparingly provided with brood-food, one is tempted to leave these cells in the hope "that at least something will come out". The bad habit of regrafting a badly accepted batch on the following day is widespread. This practice should be rejected on principle; firstly, there would be pupae of different ages in the batch and, secondly, the quality of the queens produced by the regrafting may very well be worse. It will indeed sometimes succeed in teasing out an extra queen from the colony, but because of inad- equate feeding she would show miserably poor development compared with her one day older sisters. If an unsatisfactory nursing mood is observed (recognisable by poorly drawn out queen cups and meagre provisioning of the cells (Figures 1,2, page 15) when a check is made on the day after grafting), it is consequently much better to discard the whole batch. The nursing colony must then be reorganised or another must be prepared and a new batch given to it. If the intention is to raise a large number of queens, then from the outset, more nursing colonies should be prepared than are actually needed.

15 From the results of the first batch a decision can then be made which colonies should continue as nursing colonies for the subsequent batches and which colonies should be transferred from rearing to some other use. The best preparation for nursing colonies begins the previous Autumn with appropriate steps (stimulative feeding, strengthening). Overwintered "super stocks" will give the best nursing performance. It should be obvious that all nursing colonies should have been brought up to the optimum condition by stimulative feeding, moving, strengthening or expansion, before insertion of grafts is made. This applies particularly to early breeding, which usually takes place at a time when colonies have not reached "breeding ripeness" without additional encouragement. The necessary procedures will depend on the prevailing conditions.

Grafting Every serious minded breeder, who regularly rears a great number of queens will use the grafting method. For the small apiary other methods will suffice (cut comb method; strips of cells; punched cells) (I.2.) but for continuous use they are too complicated and not very satisfactory. The grafting method is in use all over the world, and we have repeatedly shown that the fear of damaging the larva by this method, though often expressed, is completely groundless. We set down here a number of sug- gestions to make the work easier for anyone who finds lifting out the larva difficult. 1. Cut away the sides of the cell to give a better view of the larva and make removal easier. 2. Use a good light source which shines on the base of the cell. Forehead lamps such as dentists use or small halogen lamps (Osram "Minispot") have proved effective. 3. Use a lens (grafting tools fitted with a lens are now available). (1.3.) 4. Choose a good slender grafting tool (e.g. the "Swiss tool" with a bent neck, Fig. 3). 5. Feed the breeding colony, or hang the comb of eggs in the nursing colony one day before the hatching of the larvae so that the grubs "swim" in the brood-food. It is often recommended that the queen cup should be provided with a droplet of "royal jelly" and the larva laid on it. I have never detected any benefit from this - grubs that are "dry grafted" are accepted just as readily. There is however one exception; wet grafting must be practised when rearing in queen-right colonies. The larvae are not harmed by being transferred at low temperatures (as for example during days of bad weather in May). On the other hand, high temperatures can be a danger to larvae, if at the same time the relative humidity is low. Unless a man can work very quickly, the relative humidity in the grafting room must never fall below 50% (hygrometers sufficiently accurate for this purpose are readily available from suppliers of scientific instruments).

16 Queen-cups It has become clear that the diameter of artificial queen cups is quite remarkably important for acceptance and for the later development of the queens. The mandril manufactured according to Zander's instructions has a diameter of 8mm. By measuring the diameter of naturally built queen cells, it can be convincingly shown that at the widest part (the bulge) the diameter is in fact 9.5mm and lower down it reduces to 9mm. We have used a diameter of 9mm for our wooden mandrils for many years, and with this small alteration, which is now largely accepted, in practice we were able to obtain not only better acceptance, but also bigger queens. Dr. Weiss has been able to confirm these findings by meticulous research. (Zschrü für Bienenforschung 8. 1968 pp. 130.145). Consequently it must be made clear, that all methods which leave breeding material in narrow worker cells (cell punching, strips of comb with eggs) yield less satisfactory results than grafting into wider cups. The factor "width of the cell cup" is clearly just as important as the age of the larvae (within an upper limit of two days' at the start of queen rearing.

17 The same good results are also obtained by re-using drawn out cells cut down to about 10mm, and cleaned, after soaking, to remove all trace of the cocoon and old food. The wax used (virgin wax, or old wax) has no influence on acceptance. Natural queen cups on old comb are normally made from old wax. Plastic cups have recently been introduced and give good results. The frequently practised 'familiarisation' of the cups in the nursing stock prior to grafting serves no useful purpose. (Dr. Weiss).

The Age of Larvae; Queen Rearing from the Egg On theoretical grounds the conclusion must be reached that larvae which receive queen-nursing as soon as they hatch from the egg produce particularly well developed and productive queens, as is assumed in the case of swarm and supersedure queens. It is by no means easy to make sure that queen nursing takes place from the very moment of hatching. Usually the bees pay no attention to eggs till they hatch. When this takes place, a queen-cup will be built round the larva, and only then will it receive a supply of royal jelly from the nurse bees. Dr Z. Örösi-Pal was the first to describe a method suitable for practical use; as with double grafting, a normal rearing is started by grafting into a wax queen cup; the next day the larva is removed and replaced with a punched egg together with its cell base. The larva which then emerges will immediately be tended as a queen larva. The double provisioning of the queen cup is expensive in labour. Accordingly, Dr. Karl Weiss (Erlangen) has devised a method which makes do with a single operation as when rearing from larvae: Groups of cells (not single cells) are cut from a virgin comb where the queen has just laid, and fixed in a rearing frame with the mouth of the cells facing downwards in the usual way. In each group only one egg is retained. When single cells were used either the workers removed the egg or queen nursing only began some considerable time after the larvae had emerged, but by the Weiss method queen rearing began by the construction

Figure 4. Worker larvae of precisely known age (the figures indicate age in hours). The queen was restricted to an area of comb for 2 hours; 3 days later the larvae had hatched from the eggs laid there. Their age is therefore known to within 2 hours. The figures give the upper limit in each case. 6 therefore implies "between 4 and 6 hours old". Immediately after hatching the tiny transparent larva lies in a minute drop of fluid. 4 hours later the larva has scarcely grown but the drop of fluid is obviously bigger. At 18-20 hours the larva is now very clearly visible, the larval food almost covers the cell base, this is a very convenient stage for lifting the larva out easily together with a drop of larval food. One day old larvae 24-30 hours, are recognisable by the greatly increased curvature of the body. This is about the limit at which they should be utilised. Especially from this stage increase in size takes place (48 hours); these larvae are now too big for grafting.

18 19 of a queen-cup even during the egg stage. This therefore permits technical breeding from the egg at a wholly acceptable cost. Queens raised from the egg are as big and equipped with ovaries as well developed as those of swarm queens. But do they, in consequence, produce stronger stocks and higher yields? This problem was studied intensively at Erlangen for three test periods (each of two years duration). The result was a definite "No" to this question. In no year were the "Egg queens" better than those raised by the usual methods. So it is right to conclude that breeding from the egg is not justified in view of the greater expense, and is not likely to attain any practical significance. The same can be said of double grafting (in which the procedure is as follows:- Graft with larvae without regard to age; these are removed after 14- 24 hours, and suitable (young) larvae from the selected breeding stocks are grafted into cells already supplied with food and on which extension has begun). The usual method, which requires less work, will, if proper care is taken, produce queens which are just as well developed and productive.

Selection of the Larvae How careful must we be in the choice of larvae for grafting? As everyone knows, bees are not choosy when the choice is left to them, as in the common practice of making queenless nuclei which must raise queens for themselves (a most unsatisfactory breeding method). These queens are usually small and most of them emerge after 10 days - sure sign that queen rearing started from older larvae. Zander and Becker demonstrated by experiments over 50 years ago that queens reared from larvae that are more than two days old do not attain normal size or the complete number of ovarioles . Many research workers, and most recently Dr. K. Weiss, have applied themselves to the problem: What is the ideal age for larvae at the moment of transfer, if they are to develop into the highest quality queens? It has become clear, using larvae ranging in age from very young (only a few hours) up to 1 1/2 days, there is no measurable difference in the development of the queens. Only when larvae were grafted at more than two days old was the size of the queen and also the number of ovarioles reduced. In practice this means that no extreme care is needed in the choices of larvae so long as the maximum age is set at 1 1/2 days. Normal practice should be to select larvae of 24 hours old or less (Figure 4). This gives a small working margin, when for example, owing to bad weather some larvae are not nursed immediately. Between the ages of 1 1/2 and 2 days a very distinct stage of development takes place. The age of the larvae represented in Figure 4 is dated accurately (± one hour) and can serve as a standard of comparison. With some experience it is not difficult to select larvae of the right age. From this illustration it can be clearly seen that larvae between 18 and 24 hours old are already well provided with larval food and are big enough to be picked up and transferred quickly and without risk of injury.

The provision of larval food is critical in the development of the queen. If a great surplus is continuously available during the later stages of growth, then the larvae can apparently make up for any retarded development during the first 48 hours without any disability remaining. In the cell of a well nourished queen there always remains an appreciable residue of unconsumed larval food. If this partly dried up food reserve is not present it is usually a sign of poor nursing. The disc of food in the open cell (Figure 2) is therefore an indicator of the nursing-zeal of the stock. The size of the cell and the speed of its construction (Figure 5) likewise afford some indication but this is not always reliable: not infrequently one finds that quite small meagre queens have emerged from large well proportioned cells.

Breeding Systems There are innumerable breeding systems but all are based on the following biological presuppositions:- 1. The bees of the rearing stock must be in an emergency mood. To bring this into being, contact with their own queen must be broken (dequeening) or impeded (rearing in a queen-right stock). 2. Presence of an abundance of nurse bees with developed food glands. Point 2 is taken into consideration when choosing the nursing stock (a strong colony in active brood production). In order to attract nurse bees to

the breeding cells, it is often recommended that combs with open brood be placed on each side of the frame with queen cells. On their own the grafted cells would be in a biologically unnatural position - isolated in an empty space in the middle of the brood nest which at the start would attract building bees rather than nurses. The use of young brood as a "mag- net" for the nurse bees is therefore quite proper, but the method has the drawback that the beekeeper must watch out for the construction of "wild" queen cells on the brood combs. The "emergency mood" is absolutely necessary for the start of queen nursing and therefore for the start of breeding. Once the queen cups are drawn out and provided with foodstuff, the queen nursing will continue, even with the queen still present so long as she is hindered from direct access to the cells. Since, moreover, the queen cells are sealed 4-5 days after grafting and need no further attention than the maintenance of constant temperature and humidity till the queens emerge, it would be foolish to use a queenless colony for this phase of the rearing. Queenless colonies work badly, many bees "desert" to nearby hives, and the rest fail to make use of their potential energy for brood rearing and foraging. Accordingly, it is advisable to carry out rearing in two successive stages. The breeding is started in a queenless colony, or in the queenless half (containing brood) of a stock which is "biologically" separated from the

22 queen-right half. The nursing colony for this stage of the process is designated the "starter". The breeder grafts will remain in the "starter" only as long as it takes to make sure they are being properly nursed and fed, which will generally be the case after 48 hours. Under favourable conditions 24 hours or even less may suffice. Then this nursing stock is ready to receive another batch of grafts or it may be used as a normal honey producing colony. The breeding batch which has been tended for 48 hours is placed in the honey chamber of any suitable stock (the "brooder"). The honey chamber (I.4.) must be fully occupied by bees and a number of combs with open brood and nurse bees placed on each side of the queen-rearing frame with the started cells. Obviously an effective queen excluder must be placed between the brood chamber and the honey chamber to make certain that the queen cannot get near the batch. Experience over decades has taught that queens are tended just as well in the end in a honey chamber as in a queenless colony. Queen rearing using this method proves to be substantially more economical. Generally speaking the methods for the last stage of rearing are broadly the same, but for the start there are several possibilities all tested and recommended. Among the available rearing methods there are a few which provide opportunities to take advantage of special situations, such as rearing in a dequeened swarm, (and also in an artificial swarm) or in a "collec- tive nucleus" (that is, a nucleus made up of brood combs from several hives). One of these methods will be used when swarms or surplus brood combs are immediately available. For rearing according to a predetermined schedule three methods are generally enough:-

1. Rearing in a Dequeened Colony For carrying out what is on the whole the commonest procedure we have a choice of two methods. Dequeening by caging the queen during the preceding nine days, or removal of the queen immediately before the rearing begins. (The quick dequeening of the colony). The difference between the two methods is that by the former no open brood will be present, whereas by the latter the nursing stock will naturally contain brood in all stages. Both methods require a search for queen cells before the first batch is inserted; with the "quick dequeening" method the search must be repeated before another batch is inserted. The method with the 9 day caging of the queen is troublesome, and often not sufficiently adaptable. Accordingly most practitioners do not cage, but insert the batch of grafts a few hours after dequeening.

2. Rearing in a Temporarily Dequeened Colony This procedure is a combination of methods 1 and 3. A start is made in a queenless colony. Early next day the queen together with the brood box is returned and the rest of the nursing follows in a queen-right colony over

23 an excluder. By this means an ample and reliable acceptance can be expected, but in addition, since the nursing stock only suffers a temporary disturbance it works from then on like any normal honey gathering colony in the apiary. So this method can be recommended to beekeepers who require only a few queens and who therefore wish to incur the least possible expense. For anyone who works with a storey-hive (I.5.) the preparation for rearing is a very simple and speedy affair.

3. Rearing in a Queen-right Colony For all the methods it is most important that only really strong colonies are used brimming over with bees, which fully occupy two brood boxes. (Figure 6). Anyone who thinks that he must economise on bees, and cannot sacrifice a stock for breeding purposes, must certainly not be misled into selecting a defective stock for nursing. He cannot expect by such ill- advised parsimony to produce queens capable of building up splendid colonies. By contrast, there are methods by which a strong productive stock can be used for nursing without perceptibly reducing the honey yield; either by breeding according to Method 2 (temporary dequeening) or by rearing in a queen-right stock from the start. This procedure makes use of the requeening instinct of a colony; if the queen has only slight contact with one part of a stock, this part will feel itself queenless and be inclined to nurse queen cells. For this purpose, it is enough to separate part of the hive by means of a division board with a small piece of queen excluder in it. As with natural supersedure the number of cells tended is fairly small (8-15) but the queens so produced are exceptionally large and vigorous. This procedure is suitable for protracted rearing by a beekeeper who needs from time to time only a few queens in a batch (e.g. for artificial insemination). The acceptance of larvae is usually poorer for the first batch than for those that follow, especially in Spring. It seems as though the bees in a queen-right stock must first "learn" the business of nursing. Consequently it is often advantageous to begin this procedure with Method 1. The first batch of the season is placed in a queenless colony. After 24 hours it will be transferred to the rearing part of a queen-right colony which had been prepared as described for Method 3. When the cells are sealed, they can go to the honey chamber of another colony and the second and later batches can be introduced directly into the queen-right colony. For both methods of starting, in queenless or queen-right colonies, it is important to pay attention to what was said above about the nursing colony (page 13). To get a "full house" in the dequeened colony, the honey chamber can be removed and the bees compressed into one box (Figure 7). In the queen-right stock the appropriate action is to restrict the space till all combs are thickly covered.

In what follows these procedures are set out in note form.

24 I. Rearing in a queenless colony.

1. Remove the queen (alone or if from a very strong co.ony with two brood combs to form a nucleus; strengthened with combs 'bled' from other colonies it may form a new nursing colony within four weeks). In the middle of the brood nest leave space for a single comb. Remove and dispose of the honey chamber. If there is not room in the brood box for all the combs with brood, combs with mainly open. brood (but without bees) are distributed to other stocks. If the hive is in a beehouse, place screens on each side of the entrance to discourage a mass migration of the queenless bees. 2. 2 to 3 hours later- Insert the grafted breeding frame. Not more than 25 grafted cups. Continuous feeding, with a bucket feeder with a reduced opening (about an inch in diameter). Daily food supply about 1/2 litre. However, many breeders prefer a heavy feed immediately after the batch is inserted. 3. 48 hours later- Take from the brood chamber of a strong productive colony four combs of open brood (Take care ! without the queen. If by chance the queen cannot be found, the combs should be brushed free of bees to make

25 quite certain).'These combs are placed in the honey chamber. In the middle of them leave a gap into which the comb of grafts from the starter colony is immediately placed. The nurse bees clinging to the frame of grafts can mainly be left to join the "brooder" colony. In this way uninterrupted nursing is ensured. The starter colony must be provided with a similar reinforcement of young bees so that it can endure repeated "blood letting" of this magni- tude. The second batch is now placed in the nursing colony. 4. Another 48 hours later (therefore 4 days after dequeening); Remove the second breeding batch to the honey chamber of a new brooder colony, between brood combs as in the case of the first batch. Place batch 3 in the queenless nursing stock. Continue feeding in small quantities when there is no nectar flow. 5. 11 days later (15 days after putting in the first batch) the 3rd batch is ready for caging. The nursing stock is brushed to provide the mating nuclei (p.36) One ripe queen cell will be left for the bees remaining in the hive.

II. Rearing in a Temporarily Dequeened Stock 1. Close to the chosen (very strong!) nursing colony, place a floorboard with the entrance towards the rear. 2. Lift off the honey chamber. Transfer the brood chamber with the queen to the new floorboard. The honey chamber replaces the brood chamber on the original site, with the flight direction unchanged. 3. Take four combs with sealed brood (or with some open brood) with bees (no queen!) from the brood chamber and transfer them to the honey chamber leaving a free space between them for the frame of grafts. 4. 3 to 4 hours later: Insert the grafts. 5. After 24 hours re-unite the colony. The brood nest with the queen returns to its old position, the honey chamber with the grafts is placed above a queen excluder.

III. Rearing in a Queen-right Colony 1. Preparation of the hive:- A "long idea" hive is required of 20 comb capacity. If the hive is established in a bee house, the frames will be arranged "warm way"; if in the open the "cold way" can be used. Put on a cover board, or a quilt and insulation. Excluder: A vertical division board, which fits closely to the walls and floor; with a window 15 x 8 cms in its upper third, covered with a piece of queen excluder. Breeding frame with a food trough: The food trough is formed by fixing a cross bar about 7cm from the top of the frame and nailing hardboard on each side of the frame, as far down as the cross bar. The rectangular trough thus formed is caulked

with hot beeswax. The bars carrying the queen cups are in the lower portion of the frame underneath the feeding trough. 2. Preparation of the nursing colony. Rearing in a queen-right colony can begin as soon as the nursing colony has 16 frames well covered with bees and an abundance of open and sealed brood. While rearing is going on it can be cautiously extended as soon as building has taken place in the vacant space. Division of the colony into two brood nests: In the flight hole section behind a sealed comb of honey and store combs (with plenty of pollen!), several brood combs with the queen, then the division board with excluder (Figure 8). Behind the excluder: combs with open brood, a space for the rearing frame, two more brood combs, store combs. 3. Introduction of the breeding material:- Grafts on royal jelly, 15 cells per batch at the most; immediately after they are put in, add food (honey syrup) to the breeder frame trough (repeat 3 hours later if necessary). 4. 2 days later: Transfer Batch 1 to an incubator colony, introduce Batch 2, and feed. 5. 4 days later: remove Batch 2, and rearrange the stock. Take 4 combs with open brood from the front and exchange for the four sealed combs

27 from the back. If necessary provide space for uninterrupted egg laying. The four combs with open brood go into the nursing section arranged as before with a space for the grafts. Introduce batch 3 and so on. Under favourable conditions 12 - 15 splendid cells per batch may be obtained. In the course of the breeding season the nursing stock will develop an urge to swarm. Therefore, the queen's wing should always be clipped! Just as soon as queen cells are reared the stock is at once dequeened and work goes on with the queenless colony. 8 days later, either a ripe queen cell or a virgin queen can be introduced. In the U.S.A. the same method is employed in the upper chamber of a storey hive. Between the lower brood box with the queen and the upper brood box containing the nurse bees, a box without brood is placed, thus effecting a separation of the two brood boxes. The batch of grafts is placed between combs of open brood in the upper chamber. Here "conveyer belt" breeding is practised. Every 4 days the stock is re-arranged and a new batch is inserted; sealed brood is returned to the lower brood box and 2-4 combs of open brood are brought up; the last but one batch is caged (that is after 8 days) and a new batch is inserted. Two batches are therefore being raised at the same time in one stock; when the new one is put in, the previous batch is ready to be sealed and the one before that is mature enough for caging. With the more widespread use of the storey hive in West Germany further experience of "queen right" rearing in a double brood box hive has been obtained. The conditions necessary for success are two brood boxes thoroughly crowded with bees, and favourable weather (mid May to mid July). For earlier or later rearing it is better to start in a queenless colony. In preparation for rearing in a queen-right colony in a storey hive 4-5 frames of unsealed brood are moved into the upper box, with an empty space left in the middle. The two boxes are separated by a queen excluder covering the whole area of the boxes. A few hours after these frames have been put in position a small batch (15-20) of grafted cells is placed in the waiting space between the combs. Continuous slow feeding is given above the batch of grafts. Four days later the sealed or almost sealed batch is moved to the side of the box; a new batch is introduced flanked by two combs of open brood from the lower chamber. As mentioned earlier, using a queen-right colony from the very beginning as a "brooder" has proved very effective. This implies that the colony receives, between brood combs in the honey chamber, a batch which has been given a "flying start" in a "starter" colony. Open brood and queen cells together attract nurse bees into the honey chamber: nursing activity is quickly in full swing, and after this batch has been capped a start can be made with Batch No. 1 of the freshly grafted queen cups without more ado. In Germany, queen rearing was also tried in queen-right stocks housed in Leaf hives. The honey chamber is half closed, i.e. one screen remains open, the other is covered. The batch of grafts is placed between two

28 frames of open brood at the side of the honey chamber which lies above the cover board. Satisfactory results can be obtained by this method, but only at swarming time. Nonetheless the method is mentioned here because it plays a part in the distribution of breeding material to beekeepers (see the next chapter).

Evaluation of Breeding Systems Anyone who wishes to raise a large number of queens at one time e.g. for despatching occasional parcels of queens to a Mating Station, will do better to work with queenless stocks. I admit that we reject the formerly common practice of raising great batches of queens, 60 or more at a time even by this method, but it is still possible to obtain a great number of well developed queens all at once. Moreover, at the end of the nectar flow, in August, it is often possible to go on working for late batches, longer and with satisfactory results in a queenless colony, than is possible in a queen- right colony. If however, smaller batches over a longer period are required without repeatedly dequeening, rearing in queen-right stocks is clearly the ideal. The number of queens reared can, of course, be arranged by using as many nursing colonies as may be required; 5 queen-right nursing stocks will yield every second day 60-70 queen cells, therefore somewhat more than three queenless stocks. What is particularly attractive about this method, as experience has proved, is that practically all the queens are large and admirably well developed; "ill favoured" ones rarely occur. This is no doubt the reason why a commercial breeder such as Piana in Bologna uses this method exclusively in his business.

Dealing with Ripe Queen Cells We strongly recommend the very late caging of queen cells - on the 11th day - since they can find no better conditions than in the colony. Here no "drop outs" or other mishaps can occur, and moreover, the humidity is always correct. On the last day before emergence the cells are scarcely affected by vibration. When there is a very good nectar flow it is not unusual for the bees to build over the sealed cells (Figure 9) often so thoroughly that the exact position of the cells cannot readily be identified. If they are examined against a window, they stand out as dark shadows. At the very least, the tip of the cell must be freed by paring away the comb with great care, so that the queen can emerge freely. If the cells are to be utilised immediately by introduction into nuclei or mating colonies they are best carried as Guido Sklenar taught us: laid in a flat box on soft padding, and held with the tips downwards under one's shirt next to the bare skin. A more modern method is to place the cells in holes bored into a block of rigid foam plastic. They will tolerate a journey of many hours duration so long as shaking is avoided.

29 After introduction into a nucleus the cells must at once be tended by the bees. Therefore they should not be introduced into a freshly made nucleus, or into one just dequeened, till a few hours have passed. If one wishes to check the contents of a ripe cell (age and condition of the queen) it can be opened without any danger to her. It is done by taking advantage of the fact that the cocoon, which is difficult to cut, only lines the lower half of the cell adjacent to the tip, while the base is made of wax only. A cut round the base allows the cell to be folded back, and the contents examined (Figure 10). Afterwards, the pliable wax will make a seam- less closure of the cut cell.

Emerging Cages In place of the wooden emerging cages formerly in use, hair curlers with an inside diameter of about 15mm obtainable at any chemist's shop are now generally employed. A wooden block with holes of suitable size holds them in place. (Figure 11). At the bottom of the hole is placed a little candied honey about the size of a cherry stone as the first provision for the queen. As with other emerging cages the queen together with attendant bees can remain here undisturbed for a few hours; with an increased supply of food she can remain for 2-3 days. For queens emerging in an incubator only thoroughly reliable apparatus should be used. No great expenditure is needed for this; a carbon filament lamp and a thermostat together with an electrical relay (obtainable from Messrs Jumo, Fulda) mounted in an old refrigerator serves excellently for this purpose. In addition to a constant temperature of 35 degrees C a con-

stant high humidity of 60-70% is crucial; this can be readily obtained by inserting a shallow tray filled with water. The provision of a hygrometer for continuous control of humidity pays off very quickly. If the humidity is kept constant at a very high level, good emergence results can be obtained by caging soon after the cells are sealed. The incubator must be checked at least twice daily, morning and evening. Emerged queens must be provided at once with some young bees and a fresh piece of candied honey the size of a cherry stone. They should not be left in the small emerging cages longer than 24 hours however.

31 Distribution of the Breeding Material Since it has been clearly shown that district Mating Stations are only of value in combination with a surrounding pure breeding zone, the conver- sion of a large number of colonies in a particular district has acquired unusual importance at the present day - as also has the question of beekeepers who are not breeders. Linked with this is the problem of a cheap, labour saving and effective introduction of a large number of queens. Generally, queens, either mated or unmated, are distributed. The first suggestion, i.e. mated queens, is expensive and therefore rarely possible on an adequate scale. Without knowledge of the proper techniques, the second alternative (distribution of unmated queens) often results in a high rate of failure. Nevertheless, if the aim is to requeen in a restricted area, the use of pure-bred mated queens should be considered, since this procedure is likely to be the most effective. However, for distribution of breeding material on a large scale, it is possible to make use of different stages of development. Transport of larvae in swarm boxes even over considerable distances, is well known and tested, but there are simpler methods which do not require nurse bees. To obtain good results, the exact stages must be chosen which are the least susceptible to deprivation of warmth and food, as well as to the effects of vibration. Dr. Weiss has tested the different stages of development with regard to this in most searching investigations. He comes to the following con- clusions:-

1. Eggs Young (newly laid) eggs (up to 1.1/2 days) are very sensitive to chilling, but older eggs can withstand a stay outside the colony of up to 2 days very well If a parcel of eggs is to be sent away, the queen must be caged on the comb for 24 hours and then the comb must remain in the colony for another 36 hours before despatch. The youngest eggs are then 1-1.1/2 days old, the oldest 2.1/2 days. Changes of temperature and humidity within the normal range have no effect. In order to develop, the eggs must be placed in the centre of the brood nest of a strong stock. The despatch of eggs by post can be quite successful when eggs of this age, round about 2 days, are used. The piece of comb with eggs must be carefully wrapped in plastic sheet (beware of drying out!) and packed in a box with soft padding.

2. Larvae and Pupae Young larvae up to about 1 day old survive for 24 hours away from their colony at room temperature without many casualties, but high humidity (over 60%) is essential. Larvae from 1 to 3 days old fare much worse and the survival rate does not begin to rise again till the last day before emergence. From these data two methods of transport and rearing can be recommended:-

32 A) . Transport of Grafted Queen Cups without Bees (Breeders' Group Schönung, Mannheim) A row of holes 15mm in diameter is bored into a block of rigid polystyrene foam with a sharpened brass or copper tube. The diameter allows the queen cup holder to fit firmly in the hole with the cup downwards. A clearance of not more than 5mm is provided between the edge of the cup and the bottom of the hole. The queen cups are dry grafted at the breeding apiary, inserted in the polystyrene block and taken to the beekeeper's apiary. Larvae have survived three hours outside the colony without suffering any perceptible losses. The beekeeper's stocks will have been got ready by moving 3-4 brood combs (with plenty of open brood) into the honey chamber. The grafted queen cups at the rate of 2 per hive are attached to brood combs in the honey chamber. 10 days later the beekeeper removes the brood combs together with a sealed queen cell and forms a nucleus for it. In this way an inexperienced beekeeper can be supplied with breeding material. This method is obviously simpler than the transport of larvae in swarm boxes. It can only be relied on to succeed during the swarming season.

B) Transport of Queen Cells The despatch of young virgin queens is easy enough, but allowance has to be made for a high rate of failure in the introduction into nuclei or mating nuclei with brood. By contrast, there is no problem with the introduction of queen cells. The despatch of queen cells even over great distances is possible if young recently sealed queen cells only are used. In these cells the pre-pupa is just spinning her cocoon, she is very lively in her movements and is apparently insensitive to vibration. We once shipped more than 200 cells in breeding frames express from the Allgaü to Oberursel (about 200 miles as the crow flies) practically without damage. The cells were accommodated in a colony to keep them warm, but they could probably have been sent even without bees. A second even more suitable stage for despatch, is the ripe queen cell, just before emergence. Hundreds of ripe queen cells have been carried in "cold bags" warmed to 35 degrees C for more than 12 hours with too few failures to be worth mentioning.

Mating Nuclei

The Nucleus Boxes. 1. The Standard EWK (Einwabenkästchen = One comb box). This design was developed by Zander, and discussion about it is unnecessary. The EWK has been tested many thousand times and has successfully resisted all attempts at modification. For many purposes it is indispensable (transport over long distances, big Mating Stations). Filling the box with exactly the right amount of young bees for its capacity is quite critical. (Figure 12).

However, the system has its drawbacks:- the (unbiological) absence of an alley way between combs (heat regulation); the small area of comb (which limits the queen's egg laying); complicated removal of the queen (particularly troublesome with artificial insemination); labour expensive cleaning at the end of the season (glass sides). For these reasons the old DWK (three comb box) when made in modern materials is enjoying increasing popularity.

2. The Kirchainer Polystyrene Mating Box With its good insulation, three small combs and large food chamber, this box supplies excellently the biological needs of a miniature colony. (Figures 13 and 14). A queen can be left in it for weeks on end and till late Autumn; the nucleus can be requeened again and again. With its lightness, ease of transportation over short distances, and its open arrangement - with no frames - it is simple and convenient for the beekeeper to operate. The quantity of bees required is only slightly larger than for the EWK. (Figure 14). The Austrian mating box is of similar design. Since their introduction more than ten years ago these multi-comb polystyrene boxes have given outstanding proof of their value under the most difficult conditions. This is quite the best type of mating box at present available, and is to be recommended wherever its use does not contra- vene particular regulations (e.g. Mating Station rules). Gradually the conviction is gaining ground that the strict requirement for complete inspection of the nuclei - at least in the smaller Mating Stations - is excessive, and allowing the use of multi-comb boxes is to be commended. For in the first place, serious offences against the regulations to preserve freedom from drones are by no means ignored and there are

adequate sanctions to forestall such incidents. Secondly, the number of drones that might possibly be brought in is in any case negligible compared with the number that fly into most Mating Stations from the outside at the present time.

The Bees The breeding techniques so far described - starting and rearing in separate stocks, breeding in queen-right colonies - make some small changes in the provision of bees for the mating nuclei necessary, since there are scarcely any queenless nursing stocks available for brushing. There are three sources for the provision of young bees for the nuclei: 1. Reserve stocks which were not used in Spring. By using them with great care - making sure that the queen's rearing ability does not suffer by taking away too many bees at one time, and by regular feeding - a reserve stock can supply an astonishing amount of bees (or brood) in the course of the season. It is recommended that the new reserve stocks for the following year be built up very early in the season. Ripe queen cells from the very first batch of grafts are used, which are placed in mating nuclei, for the

35 queens to be apiary mated. These nuclei will still get some benefit from the nectar flow and so develop into quite splendid young stocks. Anyone who has worked his way through this procedure will have little cause to complain about scarcity of bees again. 2. "Bleeding" stocks, in the process of swarm prevention. From a stock in tip-top condition more bees can be quietly removed, in the routine of swarm prevention than would be necessary for that purpose alone and without harming performance and activity. (Bees from colonies with queen cells which have already been laid in are not suitable however, since the acceptance of virgins is poor.) 3. Bees from colonies which were not ready for the nectar flow in time, provided that there is no disease. In all three methods the bees are of better quality, better than those - some of which are too old - taken from queenless nursing stocks. It is taken for granted that in the artificial swarm which has been produced there is a preponderance of young bees. When "Bleeding" a stock, we use as a rule the Marburg Separating Box. (I.6.) which ensures that all the old bees fly away. It also makes for speedy working since it obviates the need for a long search if the queen is not found at the first attempt. With this box, even in cold weather most of the flying bees return to the colony instead of going through the screen. Even if a stock is thoroughly brushed, bees will not be removed completely from the hive sides and the outer combs. These remaining bees, together with the flying bees that return and some of the brood, will result in a small stock which when provided with a queen cell from the batch will develop into a good reserve colony. Two rearing methods have been described quite intentionally, in which the nursing stocks are not broken up, but survive. Furthermore, by repeatedly starting in a queenless stock and finishing the nursing in the honey chamber of a queen-right "brooder" stock, appreciably more queens will be raised than could be accommodated in nuclei made up from the bees of the nursing stocks. The provision of bees for making up the first nucleus boxes has just been described. With the increasing use of multiple comb mating nuclei the possibility of using the same mating nuclei time after time also increases. In countries with a longer mating season and good uninterrupted mating weather, as for example in Italy, the mating colonies can be used up to ten times. But in our country a mating nucleus can only be used up to three times, of course only on condition that it was strong to start with, and that neither of the first two queens was lost, but laid well in the brood nest. As soon as the first lot of brood is sealed the young mated queen is removed. 24 hours later, a ripe queen cell can be introduced. (With immediate introduction acceptance is uncertain). Mating nuclei in Kirchainer boxes in conjunction with the breeding procedure described above make possible the especially "cheap" rearing of a greater number of mated queens. For this the following method is proposed:- The first (early) breeding batch is placed in a temporarily

dequeened stock. With the resulting 25 queens, 25 Kirchainer nuclei are made up. Two to three weeks later rearing is started in the honey chambers of two very strong queen-right stocks. Meanwhile 20 young queens have been mated and have commenced egg laying. As soon as the queen cells of the second batch are ripe the mated queens are removed for use and 24 hours later they are replaced by the ripe queen cells. Result - 15 mated queens. The same procedure is repeated 3 weeks later; from this late breeding 10 queens can probably be expected. (The queenless nuclei could be re-used only in part). The final result is therefore 45 mated queens; for which only the expenditure of a substantial quantity of shaken bees for making up to 25 nuclei is to be charged. Such considerations are relevant, for in queen rearing too, the principle of economy is to be observed.

Filling the Nucleus Boxes A sufficient quantity of young bees must first be obtained and brought to a state of quiet with food in a cellar. When required for filling the nucleus boxes, the bees are thoroughly wetted. (I.7.) A square ladle with a 1/4 litre (or 300 cc) capacity has proved most suitable for transferring the bees; it reaches right into the corners. This style of ladle can sometimes be obtained from kitchen suppliers - it's a question of fashion. Otherwise it will be necessary to have one made specially. (Figure 15). Take

37 note of the quantity of bees; a level ladleful for the EWK, a heaped ladleful for the Kirchainer. As a rule the queen is simply run in through the flight hole. To reconcile her disposition to the bees she is previously briefly immersed in luke warm water (Figure 16). Only if queen and bees do not belong to the same race, or if many flying bees are present among the shaken bees (because of bad weather at the time it was made up), do I advise that she should be introduced in a cage with a closure plug of soft candy. Only in an emergency should the rule of a 3 day rest in a cellar at a temperature of about 15 degrees C be set aside. If the nuclei are to be sent away afterwards (e.g. to an island Mating Station) it is strongly advised that the flight hole be opened in the evening before despatch. The bees can then empty their bowels and during the night following they will substantially extend their comb building. The breeder can examine the nuclei and firmly reject all defective ones. The better end result will amply repay these small labours.

Introduction Methods The last stage in the long journey to a pure bred stock is the introduction of the queen. Countless methods have been extolled as "highly recom-

mended" or "absolutely certain" but no single method is perfect. In practice the chances of success in the prevailing conditions must always be weighed against the necessary outlay of time and attention. An important factor in achieving success is the precise time of re-queening. Before the solstice many things are permissible, which later can be guaranteed to turn out badly. But often unfortunately we cannot choose the time; we must do the best we can in our short breeding season. The most unfavourable time is in the weeks after the end of the nectar flow. (End of July - August). Later, at the end of brood rearing and after, the difficulties are again much less. There are special problems when introducing after a long journey (interruption of breeding) and with queens of a different race. Here we list in summary form the most recommended methods based on the foregoing considerations.

1. Simple Introduction Methods in the Early Part of the Season a) Rapid introduction with the Wohlgemuth cage, b) Rapid re-queening with a "one hand" cage which is simply clipped between 2 brood combs after removal of the old queen. It is recommended that after a few hours of secure enclosure the behaviour of the bees on the outside of the cage should be observed before the queen is finally released.

2. Introduction Methods for Normal Conditions during the Whole Season a) Queen under a press-on cage (made of wire not plastic!) over emerging brood. Long detention of the queen under the screen is undesirable and will be avoided if there is an "escape tube" 40mm long 10 mm diameter, soldered on at one corner and filled with candy (Figure 17). b) Introduction with the Miller cage. The principle is the same as of the press-on cage. The queen has restricted contact with the bees before she leaves the cage. In the Miller

Figure 18 Miller introduction cage. The shorter entrance tube is restricted by a piece of excluder. A hair curler can also be used as a cage.

cage this is achieved by having two entrances of different lengths and 8.5mm in diameter, blocked with candy. The shorter, about 20mm long is, naturally released more quickly, but inside, it is closed by a piece of excluder through which bees can enter but the queen cannot get out. The longer entrance, about 30mm takes another 24 hours to clear, and the queen now acclimatised can leave the cage. The Miller cage is obtainable commercially in the USA but a substitute can be made easily and cheaply from a hair curler, open at both ends, with appropriate cork or wood stoppers (Figure 18). (I.8.) c) Dequeen the colony, After at least 4, better still after 7 days requeen, by simply putting the queen on a comb. It is not necessary to break down started queen cells. But an essential condition is: the queen must be in full lay, (therefore not after arriving by post!) If the queen has been caged before being introduced (e.g. for transport) she should be brought into lay in an EWK before being placed in her new stock, prior to using this method. d) Introduction into a nucleus without open brood and without flying bees or into an artificial swarm. e) Introduction by uniting the mating nucleus with the colony (newspaper method).

3. Re-Queening under Difficult Conditions (For example a queen of a different race). Nucleus with emerging brood, provisioned with honey and pollen but no bees. To maintain the correct brood temperature the nucleus is placed in

40 the honey chamber of an occupied hive or in an incubator. Free flight is cautiously permitted only after 7 days. Attend to ventilation and water! If the necessary care for nursing the colony and guarding against robbing is provided there is scarcely any failure in difficult cases. (1.9.)

2. Rearing of Drones In controlled breeding the production of drones at the right time is more difficult than of the queens. The reasons are:- the time they take to develop to sexual maturity is the longest; rearing is very dependent on the season; and such very large numbers are required. Anyone who is able to influence the paternal side in a breeding programme (operating Mating Stations; providing a commercial instrumental insemination service) must therefore incorporate drone breeding very carefully into his programme. The inclination to breed drones varies greatly from stock to stock just as does the inclination to rear queens (Figure 19). Really good stocks, ones which are good honey producers and are also reluctant to swarm, can only with difficulty be induced to rear a larger number of drones and then only for a short time. Since there are no reliable expedients ("wrinkles") to deal with this difficulty, the breeding plan should already have been drawn up

41 in the previous year, and a group of apiary mated queens from the breeding line it is intended to use should have been prepared in readiness. These queens will produce pure drones and - since hybrid stocks are used - in whatever numbers required, and practically at the time required. Since such stocks can be expected to bring in a good harvest there should be no risk to the breeder on account of these preparations. Breeder drones must be accompanied by a scientifically attested report on physical characters appropriate to their race standards. If one year old queens are employed for drone production - which is quite permissible, since for the drone producing colonies themselves no performance records are required - then there will probably be difficulty in producing a certificate on time. In this case it is advisable to proceed as follows:- Every stock that is to be considered for use as a drone producer receives into the brood nest, even before feeding begins, a frame with patches of drone cells, or a cut out piece of drone comb. By the second half of April with the normal course of Spring weather sufficient drones should have emerged to permit an assessment of physical characters. At the time planned (breeding Timetable p. 44) a whole drone comb is given for the queen to fill. While these drones are developing, the assessment of characters can be carried out by the breeder, or he can apply to the Institute for the assessment to be made and the certificate issued. For the early rearing work, a compartment of the comb cupboard is reserved for first class drone combs. Stocks that could be earmarked with confidence would have already received a whole drone comb into the winter clustering space the previous Autumn. Once the building urge has awakened, there are generally no problems as regards filling the comb. Nothing draws the queen with such magnetic force as freshly built comb. It is obvious that in this case the frame must be fully drawn out. Often very many drones are needed from a single queen. In this case the breeding mother is only employed for laying while rearing follows in drone-free nurse colonies, either above or below the excluder. When this is done, there is one thing which must certainly be kept in mind; drone eggs are liable to be destroyed after transfer, while larvae are more likely to be accepted for nursing. In consequence, there should be a delay of four days after the eggs are laid, till the larvae hatch. The combs are then placed between open brood in the nursing stock. The best timetable can be ruined by persistent recurrences of cold spells in the first half of May. Only by careful tending (daily feeding) will the retention of the drone brood be ensured in such conditions. Provided that the breeding stock is strong there is little danger to sealed brood. As soon as flying drones are observed from any stock in the apiary the drone - providing colonies must be isolated by an excluder (see page 80). The drones begin to fly 8-10 days after emerging. They can be confined for about a week in the hive without their coming to any harm. If after all they cannot now be used then the colonies are removed to an out-apiary to one side of the main apiary. In our experience with a separation of barely 50 metres

there is no need to expect a return to the original site especially if the flight direction of the hive is altered. The drone's average expectation of life is relatively short, about 25 days, which implies only 12 days of sexual maturity. So there must be continuous provision for an undiminished replacement of drones. How the reliable supply of drones can be ensured till late Summer is described in the section Mating Station Maintenance. P. 87.

43 Live drones can only be transported in stocks or nuclei, never in cages. Insemination laboratories enclose drone semen in glass capillaries and send it half way round the world. In this condition it is kept alive for 10- 14 days.

3. Co-Ordination of Drone and Queen Rearing Early breeding is desirable both for technical management and for complete mating control. By "early" however, we do not imply extreme easi- ness, such as the time of Cherry Blossom - which might be possible, but hardly economic - but a breeding plan with the matings taking place in the first days of June. Experience shows that enough fine weather can be expected at this time, and the newly mated queens built up to colony strength by the middle of the nectar flow. The limiting factor for the start of breeding is the rearing of drones in numbers sufficient for our present day demands. In those frequent years with a cold April and May the provision of mature drones at the right time often raises serious problems. Occasionally it has been our experience to have a Mating Station stocked up with queens well before flying drones were available. Owing to the long development period of the drones exact forward planning is necessary. The accompanying timetable explains the co-ordination of drone and queen rearing. Later in the season all stocks contain drones of course, but if it so happens that an unexpected break in food supply occurs (such as could easily decimate the drones on a Mating Station in a very short time), no improvi- sation can help. We have learned by repeated experience that a complete drone comb should be left in the brood nest of all stocks which may be required to provide drones all through the Summer.

44 Chapter II Selection for Performance

1. What is Breeding? Queen breeding is in the first place, simply the increase in the number of queens. By regular rejuvenation and keeping a reserve supply of queens the output of an apiary can be very substantially increased. The enhanced productive energy of the young queens is utilised and deficiencies in the working stock, which result in queenlessness, drone-laying, or any other kind of failure in a queen, are made good. Queen breeding ranks as the most important activity in the efficient management of an apiary; by it the apiarist, in the words of Professor Schillers of Vienna, advances from being a Beekeeper to being a Beebreeder. Yet breeding is not merely a question of reproduction. Above all, breeding implies improvement in the bee's performance capability. No colony is exactly like another; brood rearing, inclination to swarm, foraging vigour, stinginess, susceptibility to disease, differ from colony to colony. Breeding means the augmentation of the best (the positive variants) and the ELIMINATION OF THE BAD (the negative variantsJ. The aim is the attainment of an apiary which is uniform and has an above average performance. Breeding is by no means a human invention. Nature, which in millions of years has brought forth this immense diversity of wonderfully adapted creatures, is the greatest breeder. It is from her that the present day breeder learnt how it must be done, excessive production and then ruthless selection, permitting only the most suitable to survive and eliminating the inferior. If this harsh rule of Nature is either set aside or relaxed the con- straints will within a short time be broken through, and a species will disintegrate into a large number of different forms which through diver- gent colouring and living requirements, through unusual body forms will depart from the natural pattern. In laboratory animals which are kept only for study and not with a definite breeding objective, these phenomena can regularly be observed. At the same time, reduced fitness for life under natural conditions very quickly results, because in nature this is only preserved by incessant and rigorous selection. If this selection is lacking because the trouble of finding food and warm shelter and taking care of the young has been removed and protection from enemies has been provided,

45 the ability to hold its own in nature, gradually but inevitably declines. What has the wolf, the ancestor of our dogs, turned into during the course of millenia under the hand of man? A Dachshund or a Poodle would soon be a pitiful wreck if it had to try to exist on its own. Nature is also the breeder's greatest instructress as to how to avoid inbreeding:- No mating within the same colony; mating flights to far distant drone assembly places; multiple mating. But breeding means not only selection from such material as is available, but also planned combinations with the aim by uniting valuable genetic material from two different lines or strains and in addition acquir- ing from the combination itself an improved performance. More will be said later about the possibility for and the limitations of this procedure as applied to honeybees.

2. The Breeding of Honeybees Our bees have certainly not yet reached the condition of the domestic animals just mentioned since even today they still have to maintain the struggle for existence largely on their own. Nonetheless, we have gone a very long way towards making their lives easier, so that the laziest colony or the latest swarm is not in danger of starving the following Winter. Today the majority of beekeepers restrict themselves to protecting their bees, but allow them to increase just as they like. What is the result? Stocks with excessive swarming tendency grow ever more numerous, while the good foragers with weak tendency to increase, which rejuvenate themselves by supersedure, are driven back. With the Heather bee, and in the Carinthian method of breeding for swarms which produce bees quite useless for a modern beekeeping business, we see the result of these practices quite clearly. At many apiaries where the increase of the so-called "Country" bee takes place without any control, this feature is plain to be seen. In addition to this lop-sided behaviour the bee is usually very stingy and excitable.

Productivity declines unless there is continuous selective breeding But the real breeder wants more. He is not content to keep his bees in their natural condition. He wants to change them, but change them in a way that suits his economic objectives. What he strives for above all is to step up the foraging efficiency well above the natural level. With bees a modest increase in performance assuredly results in substantially increased yields. As illustrated in Figure 20, it is assumed that a stock has produced a gross yield of 60Kg honey. Of this 50Kg is needed by the colony, leaving l0Kg as harvest for the beekeeper. If the gross amount can be raised by 20% to 72Kg with consumption by the bees unchanged at 50Kg there will remain 22Kg for the harvest, an increase in yield of 120%. Breeding is therefore much more than mere queen replacement, more than mere preservation of the status quo.

46 Simple queen rearing provides young, productive queens in the apiary. Breeding improves the inherited qualities of the bees. These two aims can only be combined if clearly defined principles are observed. Of course any beekeeper who engages in queen rearing will by pure intuition only breed from good queens. But this is a long way from saying that however good the progeny may be, an improvement has therefore been achieved in the heritable qualities in the apiary. Every peak performance is not by any means inheritable; the many failures with the so- called "selective breeding" (i.e. requeening from the best colony in the apiary, regardless of its ancestry) give adequate proof of this. Only too often one is deceived by a "Blender"; which certainly shows outstanding qualities itself, but whose descendants are a disappointment. Enduring success in breeding is only likely when it proceeds from breeding lines which have been tested for generations for their suitability for the forage conditions of the beekeeper, when the reliability of inheritance of their qualities has been demonstrated, and furthermore, when a breeding programme is closely followed which avoids inbreeding as far as possible and has regard at all times to the best combination of genetic material.

47 Fortunately we have a number of well tried strains, which thanks to the efforts made by conscientious beekeepers for tens of years, fully satisfy these requirements.

3. The Results of Selective Breeding Selective breeding is possible wherever heritable differences are available; and such differences are present everywhere even within "pure races" Let us just take a look at the Carnica; even in this bee there are plenty of characters and qualities which are not everywhere constant but show large variations from colony to colony. For example body colour and hair colour vary from light to dark. In some colonies some of the bees have a brown ring, others not. The Mean Cubital Index of a stock can range from over 3.00 to 2.50. Quantity of brood can be very large or small. Many colonies show an almost uncontrollable swarming tendency, while others can be managed throughout the season without showing any such inclination. There are also obvious differences within the race with regard to rapidity of Spring build- up, foraging zeal, "resourcefulness" quietness on the combs etc. Frequently, therefore there are economically important qualities which vary within wide limits, in one and the same race. The starting point for selective breeding resides in these differences. They make us aware how necessary it is to test continually, even within a "pure race" and then to separate the wheat from the chaff. Not every Carniolan is good, and the most beautiful grey coat, the longest tongue, count for nothing if in a short time the stores are all turned into brood, or the stock swarms at the wrong time. One often hears it said "We live in a Carniolan area; here all the breeding is pure, we do not need a Mating Station". Thereby one of the most important principles of breeding, the requirement for control of both maternal and paternal parentage, is neglected and any improvement through breeding made impossible from the outset. "Pure Carniolans" can certainly be bred in this way, but not the bee required for modern economic beekeeping. From his breeding strain a breeder requires above all heritable performance. He is not well served by a "fluke" or a "Blender" - for the progeny will not retain what the mother colony promised and they may perhaps be even worse than the average of the apiary. Hence, the breeding work has not benefited from it. A breeder colony which on the other hand breeds true will, with suitable mating, retain the improvement in the next generation. An apiary made up of stocks from a tested breeder-strain brings higher returns from less work, because the stocks yield more on average, they are more uniform and can be managed with less labour. Where the common "local bee", (that is to say, the heterogeneous racial mixtures in apiaries managed on some "let alone system" without thought of breeding) has been compared with breeding strains, the pure bred stocks are always superior no matter what race is involved as the accompanying table shows.

* In this experiment the pure bred stocks were a Carniolan combination (i.e. from two pure bred Carniolan strains) of the Institute at Oberursel, obtained by artificial insemination of the queens. Table 1 Comparison of some productivity results of pure bred queens, their apiary mated daughters and queens of the local country bees.

In 1967 at the Testing Apiary of the Institute for Teaching and Research in Bee Breeding at Kirchain, 4 stocks from each of 9 different strains were tested, as part of the "South Eastern Research". They were all of Carniolan extraction. As the standard for comparison the Carniolan strain of the Institute was used which for many years had been propagated at an unreli- able Mating Station and could no longer be classed as pure. The average yield of this strain in a moderate year was 8.0Kg in which individual performance varied between 4.3Kg and 10.7Kg, so the worst was only 40% of

49 the best. The most productive pure Carniolan strain (Troiseck from Lunz am See) in comparison averaged 16.3Kg, varying between 14.6Kg and 20.2Kg with the lowest reaching 70% of the highest. The best thoroughbred Carniolan strain thus easily attained to double the average with a noticeably more uniform performance. These examples could be multiplied at will. Dr. Wohlgemuth enquired of the purchasers of his queens about their performance one or two years later and found their superiority almost everywhere confirmed. The advantage in yield from a pure Carniolan strain, the Troiseck, is especially great, frequently double that of the very strongly crossed local bees of North Germany. This great advance in performance alone is sufficient to explain why the Carniolan bee has inside 20 years almost completely displaced the native German bee. Moreover, the breeder strains have achieved a 25-30% superiority in their native districts over the "wood and meadow" bees. In view of the short time during which the bee has been selectively bred, that is a great deal. But the economic advantage to the beekeeper is substantially greater than would be assumed from a casual glance at these figures. Because operating costs remain approximately the same, an increase in the yield of about 25% resulting from better breeding usually means that the "nett yield" after allowing for expenses is increased by about 50%. Today as much honey is produced in the German Federal Republic as was produced in the whole of the German Reich before 1939, in spite of the deterioration of forage, shrinkage of available areas and the decline in the number of stocks; thoroughgoing bee breeding work is undoubtedly one of the factors that has brought this about. At individual apiaries in districts where no deterioration of forage has occurred (e.g. Forest areas) a very marked increase in yield has been observed. The following production statement (from about 40 stocks) is taken from the records of the experienced Carniolan breeder, J. Dorminger (Knittelfeld). Period Average yield 1941 - 50 22Kg 100% 1951 - 60 33Kg 150% 1961 - 70 43Kg 190% The summarised results of the German and Austrian apiaries used for productivity testing and of breeders in those countries obtained over 25 years are set out on pages 77 and 78 and in Figure 30. Obviously all these data only apply to breeding lines that are suited to the forage sites and the management methods which obtain at the time. "Good" and "Bad" in an absolute sense occur but rarely in bee breeding; breeding lines with unequivocal peak performance under one management can break down completely under different conditions (p.89 and 98). In the course of bee breeding, management methods must change as the bees change. Selection for increased brood rearing makes it possible to build up stronger stocks. This in turn requires the introduction of hives that can be

extended as necessary (storey hives). In the Carnica Breeding work of the Research Station at Lunz am See the trend towards more vigorous build-up and consequently stock with enhanced production capability has been clearly apparent since 1948, that is for nearly 40 years. The bees and the system of management must be closely in tune. If a bee capable of great performance is kept in a small hive with a fixed amount of space (for example a leaf hive), the stock may need to be "bled" for swarm prevention before the nectar flow begins and will thus be deprived from the outset of the opportunity for achieving the output for which she has been bred. One thing certainly should not be overlooked:- Breeding does not so much increase the output of the single colony as the average of the entire apiary. For in apiaries where no breeding is done a "Blender" will appear from time to time and give peak results; but together with this there will always be several failures, which on the whole bring in nothing, and so considerably depress the total harvest of the apiary. At an apiary with properly bred queens, complete failures are rare, production is uniform. Consequently the total harvest (and of course the average per colony) is higher, even if the yields from the best colonies are not much different from the best colonies of the local bees (Figure 21). Therefore, one should not expect that a single bought-in pure bred queen in an apiary of 20 stocks will produce a higher yield than the best of the local bees. That would be a very unfair test. In order to find out the real difference in productivity one should rear queens from the pure bred

51 queen, and then compare the total yield from ten of her daughters with that of ten local stocks. The observations of Beemaster Krüger of Hamburg show this clearly. His Carniolan pure bred queens (mated on an Island Mating Station) produced an average yield of 114%, about 26% above the average of the apiary mated queens (which were also of a good strain, but not mated at a Queen Mating Station). Notwithstanding that, several of the best colonies with apiary mated queens brought in a harvest that was higher than the average of the pure queens. A single average pure queen in an apiary of local bees will not therefore necessarily give the maximum, but will be among the upper third of the outputs. Only the average of several such colonies will demonstrate the superiority of the pure race. Incidentally it should also be noted that the pure mated queens required 35% less attention since their uniform development allowed uniform treatment. As a rule, uniform progeny, which will breed true, will only be obtained from uniform, pure bred colonies. Starting from hybrids or from the uncontrolled mixture of local bees, many years of increase AND selection would be needed before a stable breeding line of practical use for beekeeping could be obtained. Permanent results can only be achieved in a reasonable time span WITHIN THE FRAMEWORK OF ONE OF THE NATU- RAL RACES OF HONEY REES. Hybrids can be used as honey getters but they must not be bred from. The starting point in breeding must therefore be the race, that is to say, a combination of genetic qualities, sieved and tested by Nature herself. Assessment of physical characters must be made in order to ascertain that this combination has been preserved in its original form, and has not been destroyed by crossing. Tests of performance must obviously be carried out alongside these tests. Uniformity of physical characters together with good performance warrants the expectation of uniform progeny. Pure breeding and breeding for production are not at all opposed to each other, as was often asserted in the past, but only the two together provide the basis for successful breeding. Of course breeding by itself provides no guarantee of above average progeny, not even if attention is given to reliable mating control, examination of physical characters, and selection for performance. Attention must also be given to the effect of inbreeding, which can be all the greater the more rigorously and diligently the selection is carried out. The accumulation of the best heritable qualities is of no value, if as a result of inbreeding the vigour of the stocks is destroyed and the good qualities cannot be expressed. This problem becomes particularly acute when a faultless mating control (Island Stations, Instrumental Insemination) is attained. Many a breeding line which has been bred according to what are now outdated methods (mating between close relatives, a single drone colony at the Island Mating Station), has suffered severe damage in consequence (p.63).

52 The old breeding systems, founded on erroneous principles, have frequently resulted in incorrectly equating "pure breeding" with "in-breeding" even down to the present day. A totally unjustified importance has been given to this idea by persons who wish to discredit pure breeding in favour of the allegedly hybrid breeding. A practical discussion on these matters is naturally out of place here. The technical terms for animal breeding are strictly defined, and the bee breeder too is obliged to adhere to them. (See "The Most Important Genetical — Statistical Technical terms in Animal Breeding" published for The German Association for Breeding Science by E. Ulmer, Stuttgart 1966). "Pure Breeding" means simply the mating of parent animals within a geographical race, a Breeding Strain or a Breeding Line (p.69). When it is considered that a race of bees can embrace several million colonies and a breeding strain several thousand, it is instantly clear that pure breeding in such a broad setting can readily occur without any inbreeding. Only the choice of related animals for mating leads to inbreeding irrespective of whether hybrids or pure bred animals are employed. It is therefore essential that due regard be had to the accepted defini- tions of the various concepts which relate to breeding in order to avoid unnecessary explanations (Ruttner : ADIZ 1978 pp 97-101, ADIZ 1979 p. 109).

4. The Evaluation of Performance. Colony Productivity The honey yield of a stock reflects firstly the nectar flow and secondly the productivity of the stock. It is self-evident that only colonies from the same apiary should be compared with each other. A very simple evaluation method has been arrived at over the years as a result of experience which only takes account of significant indications and is very easy to carry out. The Hive Card for Evaluation of Performance is expressly designed to suit the method and should be used by all breeders. (Obtainable from the office of D.I.B. 5307. Wachtberg 3). For carrying out performance testing at testing apiaries, general rules were worked out by an international panel commissioned by Apimondia in 1972 (Report of the symposium: Selection and Mating at Lunz am See, Bukarest 1973; obtainable from the office of D.I.B. 5307 Wachtberg 3). It is recommended that these general rules should also be considered for performance testing in private apiaries as far as possible so that the objectivity of records is guaranteed, and the comparison of results on a broader basis becomes possible. For practical use it is important to point out that the Evaluation Scale has been turned round to conform to international rules so that "excellent" is 4 and "too little" is 1. The Hive Card Printed in slightly reduced format (p. 54)

53 The Hive Card Headings should ensure that significant observations are made and they are clearly indicated with a few ticks. The lay out lets the beekeeper take in the condition of the colony at a glance during an inspection. This hive card provides every beekeeper with a convenient and complete management record; it also serves as evidence of performance in queen breeding. The queen breeder uses a separate card for each queen in each management year; in contrast the commercial beekeeper will use one card to cover the life of each queen. The particulars concerning the size of the apiary and the beekeeper's address are required at the breeding station for selection purposes, the management year is for later filing, if needed for a card index. The data on the pedigrees of the queen and the parental drone stock are transferred from the breeding card. If, however, it is a matter of natural rejuvenation then the word "swarmcell" or "supersedure cell" is marked with a tick and "Apiary Mated" is indicated at the space marked "Mating Station". The previous year's honey production and the corresponding apiary average are transferred from the previous year's cards. Similarly the first row will record the Autumn review of the previous year. (p.54) It is a matter of principle that the card travels with the queen from the nucleus to a hive, with the prime swarm into the next hive, and accompanies her on removal to another apiary. These movements are reflected in the heading "Hive No." Inserting the date ensures that a separate line is used for every manipulation. The "General Report" reflects the condition of the stock before any changes have taken place. "O.C." is the number of combs actually occupied. The number of combs occupied by brood is followed by the different growth stages of the brood. The amount of brood in each stage is assessed and points recorded in the corresponding box:- 4, excellent, 3, good, 2, low, 1, too little. From these data the seasonal changes in the arrangements of the brood nest can be seen which, as everyone knows, vary greatly. These entries have immediate use for the detection of supersedure. The beekeeper must always keep an eye on "stores" so that the bees do not go short. With a removable super it is easy to estimate the weight of honey without opening it. We thus get an idea of when the stock gathered the main harvest. At the same time, this entry (in Kg) provides a check on the extraction schedule. Gentleness' and 'quietness on the comb' must be assessed at every examination, using the same signs as for brood quality:- 4 = keeps still., 3 = restless., 2 = runs for the honey., 1 = leaves the comb. It is common knowledge that these important attributes depend only too often on the faults of the beekeeper, a particular nectar flow, or other outside influences, so that they must be assessed not just once but again and again. "Given +, Removed — ":- Empty combs, foundation, combs with brood, or bees given to, or taken from the colony are recorded here. Every kind of

55 feeding or removal of honey will be recorded in Kg of dry sugar or honey. (The card consequently allows for bucket feeding). Pieces cut out of drawn combs (-DC) come under the heading 'F.f', but if already full of drone brood, then (-DC) comes under brood combs. The size of these cut out pieces can be given as 1/2 or 3/4 of a standard comb. Entries are made as necessary in the roomy column for "Remarks" for start of brood rearing, dead bees, supering, swarm; or type of feeding etc. If the queen is seen enter ♀! If she is a marked queen, add a point ♀.! This is a check that the mother of the stock has not been replaced by a young queen. The card is started at the Autumn Review, because it is important to know for all later comparisons how strong the stock was at the start of winter. What requirements for winter food were recognised can be straight away recorded e.g. + 8Kg sugar. The amounts of food given can however be recorded later under "remarks" or marked on the hive in chalk since this is of only minor importance. The first spring records begin under the thick horizontal line. Depending on the weather it will be found if the queen is laying and above all whether food supplies are adequate. If the colony is reduced by the removal of two combs of honey this would be recorded -2C and -2Kg honey. On the other hand, if a food comb is given the record would be +1C and +lKg. If a feed of 1 litre of syrup is given it is simply recorded as half, to represent the dissolved sugar, e.g. + 1/2 Kg. If a pollen comb is supplied "P" is noted under "combs". Other incidental observations go in the ample space provided for remarks. These refer particularly to swarm preparations. Queen cups, queen cells (laid in; open; sealed) if any are found. The action taken (e.g. expansion of the brood nest, removal of the brood, or bees), will be noted in the appropriate box at the same time. All other observations of matters that interest the breeder, arrangement of the brood nest, the way in which honey is stored, use of propolis etc., can be entered under "Remarks". If in a colony there is an excessive occurrence of brown markings on the first abdominal segment of the worker bees — this can vary according to the season — it would be recorded according to the size of the spots:- small spots s; large spots S; whole ring R or RR. Possible consequences for Selection can be noted at the top of the card. This card has been designed to provide as complete and lucid a presentation as possible with little printing and little need for writing. Finally it must be emphasised that these "Hive Cards" would be better named "Queen Cards" since each is the identity card and life history of a particular queen regardless of which hive or apiary she is in. These changes of residence are recorded in the column "Hive Number" both for time and place and obviate any confusion of the cards. At the end of the year it is easy to analyse these entries and to compare the individual cards of the apiary one with another. For the commercial

56 beekeeper it is the balancing of his books; for the breeder it is the required proof of performance. The recording of behaviour and performance of individual bee colonies is not just a trivial pastime for dilettante beekeepers but is of major economic importance. Many beekeepers have all the details of every single one of their colonies in their heads, even with great numbers of stocks, but it is better to have written records in the form of hive cards, the details of which can be suited to individual needs. But it is only by the systematic collection, arrangement and use of all observations that one can get the "feel" of the apiary from the breeder's point of view. The secret of success of all great breeders — we call to mind particularly Brother Adam's results — that no essential detail concerning their stocks escapes them, and they have this information available at all times. Even for honey production pure and simple it pays to keep accurate records of the colonies. One of the most productive commercial beekeeper's in the world, Ian Stephen of Tasmania (average production about 150kg per stock), provides a convincing example. Every one of his 1,600 stocks has its own record card, on which the essentials are recorded (more particularly for the honey yield, of course). These data furnish the basic information for the selection of high performance queens specially bred to meet his requirements.

The following types of performance provide the basis for evaluation 1. Honey Production. (Harvest + stores before feeding) The yield is calculated as a percentage of the average of all the available stocks. The performance to be judged must be that achieved by a colony without outside help. Removals or additions because the colony has too many or too few bees will not be taken into account; if a colony has to be strengthened because of slow development then it is excluded from the evaluation. If it swarms, or has to be reduced as a means of swarm prevention that is its own loss. It is of very little interest what a stock would have brought in if there had been no swarm, on the contrary we want as far as possible to eliminate colonies with a pronounced swarming tendency. For this reason the yield from a swarm is not credited to its parent stock. From the yields expressed as percentages of the apiary average, we get a ranking of colonies which forms the basis for performance selection. When making a decision between colonies of similar good performance, all those which are lower down in the marking are only considered at a second review. Breeding colonies must have an above average yield. (The Breeding Regulations of the German B.K.A. (D.I.B.) require that they lie in the best 40% of at least 20 colonies) The ranking with respect to percentage yields makes possible various comparisons of colonies between different years, apiaries and breeders. It can often be shown that sister queens from the same batch are in the peak attainment groups at quite separate apiaries. This comparison is very important, for the greater the number of stocks, and the more varied the

57 environment so much better is the selection. Selection from less than 20 stocks has little meaning. For an objective comparison of productivity, the following values are determined;- Apiary average (in kg.) = Total apiary yield (in kg) ÷ Number of colonies. Colony production (as % of the apiary average) = Yield of colony ÷ apiary average x 100 Group average (in % ) = Sum of the percentage values of a group Number of stocks. Average deviation from the = Sum of the deviations from the group average group average (in %) ÷Total of all percentage values of a group x 100 The average deviation from the group average gives a measure of the uniformity of a group; the lower the value the greater the uniformity of the group. This measure can also be obtained very simply by determining the average ranking taken up by the colonies in the group in respect of their yields. For example, in an apiary of 20 colonies all are arranged in order of their yields, and each receives a place number. The numbers 1 to 10 lie in the plus-group, the numbers 11-20 lie in the minus-group. Group values can be expressed in terms of place numbers, and different apiaries can be compared with each other. 2. Spring Build-up Assessed by the time when the first super is put on. The colonies are divided into four categories ;- 4 - very early 3 - early 2 normal 1 - late (supering only just before the early summer flow, or not even then). In this matter too, assessment is made by comparing all the colonies of the same year and of the same apiary. The spring build up very often finds expression in the honey yield. Its separate assessment is specially important in a year in which because of unfavourable weather, the early flow fails. To what extent it is taken into account in breeding selection depends on the breeder's situation as regards nectar flow; that is to say whether the early spring flow plays a part in the district in question or not. In districts where there is nothing but the forest flow, a too rapid build-up can be a serious disadvantage (urge to swarm). But on the other hand it is often noticed that the colonies which have the liveliest spring build up are among the peak performers in the late flow. But combining rapid build up and great brood production with low swarming tendency is one of the most difficult tasks in selective breeding. 3. Building Urge Determined from the number of drawn out sheets of foundation and combs. 4. Number of Empty Cells Among the Brood This value gives an indication of the extent of inbreeding (see section on the Sex Alleles, p. 64). 5. Gentleness. 6. Quietness on the combs

58 For the two last named qualities there are no objective standards that can be conveniently applied in practice. Furthermore, they can vary with the environmental conditions. In order to reduce the uncertainty of these factors it is recommended that an entry be made in the hive card at every examination. At the end of the season the sum of the records will give an overall picture enabling a pretty sound estimate to be made. Gentleness and quietness on the combs are important racial characters of the Carniolan bee. If they alter from one generation to another, mis-matings can be inferred and further investigations can usually be avoided. But gentleness is also a very important character for selection in the Dark bee. Notorious stingers are not wanted either by the beekeeper himself nor by his neighbours. In thickly populated areas, beekeeping should only be carried on at the present time with exceptionally gentle bees. The continual existence of beekeeping in such areas depends on keeping bees that will tolerate distur- bances near their hives, and this quality is often more important than the size of the honey harvest.

Performance of the Family The performance of a single colony, taken by itself cannot show whether it is a chance result as a consequence of environmental conditions (a corner position, quiet robbing), a unique, one-off, favourable combination of genetic qualities ("Blender") or the result of an enduring heritable constitution. The Heritable value can only be known by assessing the whole family. For this reason the performance of ancestors and sisters must be included when making a selection. It is to be expected that the performance of the earlier generations was above the average of the apiary or they would not have been selected as breeder stocks. It is not so obvious for the sisters. Sister-testing requires that at least 6 sister queens from every generation of a breeding line be retained in full stocks (i.e. not in nuclei) in the same apiary. (At testing stations 10 test colonies from each group are required). The most valuable breeding mother is one which has undergone progeny testing. If experience has shown that the daughters have turned out well, large scale propagation can be started with confidence. It is of course taken for granted that a test batch is bred from her even during her first production year. Usually this will only be risked if the breeder line in question is known to breed true.

5. The Honeybee Family It has become customary to transfer to the bee the terms of human relationships; when referring to bee colonies we speak of mother, father, sisters, grandmother etc. This practice which is usual and permissible in other forms of animal breeding, leads to certain difficulties when applied to the bee because of the diverging proportions of the genetical inheritance.

59 A bee colony is a real family, the hive mother, the fathers (who indeed have been dead a long time, but genetically are still present as spermatozoa in the queen's spermatheca), and the worker bees which are children in the normal sense. But we must bear in mind that on average the queen mates with 6 to 10 drones and so has stored in her spermatheca as many different types of semen which in their turn carry out the fertilisation of the eggs. The crowd of worker bees in a colony is not therefore composed of full sisters, but of 6 to 10 (or even more) groups of half sisters. From the point of view of breeding for performance and character we regard a stock or colony as an animal (the"Bee") just like a cow or a hen. In reality these qualities are derived from two generations, from the colony- mother with egg laying performance and life span, and from her worker-bee daughters; in the case of hybrids the workers have in part some genetic characters quite different from the mother. The mother has a herd-book number, just like any other animal kept for breeding. To identify the colony it is customary to use the Hive Number of the apiary. If the hive number changes (because the colony has been moved to another hive, or because of swarming) a new hive number is entered on the Hive Card. However, the Queen Number remains unchanged throughout the Queen's life. The ambiguity in the nature of the colony creates difficulties again and again in the construction of a family tree. This is very clearly seen in the designation of colonies at a mating station; some speak of "Father Colonies" because they think of the drones as fathers of the future worker bees, others call them "Husband Colonies" since they provide drones to be mates for the queens. We prefer the factual name: "Drone Colonies". A diagram can illustrate bee-relationships better than words. Figure 22. Colony "A" has a queen whose herd-book number is X/80. She was mated with drones of the same strain. These drones are of course not present in person, but none the less they are effectively present as the spermatozoa in the queen's spermatheca, and are therefore shown in the diagram. The symbol for a drone is repeated three times to indicate multiple mating. The workers and drones produced by queen X/80 are all broadly speaking alike. In 1981 a queen Y/81 was reared from a larva of the Colony "A" (with queen X/80). She was mated with drones of alien origin (indicated here by empty rings) and has built up colony "B". This stock naturally contains hybrid workers, but the drones are pure bred of the same strain as Stock "A". In their relationship with workers, and any possible daughter queens that may be raised in the same stock, drones are usually described as "Drone- Brothers". As the sketch shows, this is not the full story; having developed from unfertilised eggs the drones are constitutionally like their mother queen Y/81; they are only her sons in so far as they were produced by her. Looked at genetically the relationship of drones to the worker bees of the same colony, is the same as that of the mother to her daughters. The queen Y/81 is a full sister of the worker bees in Stock "A" and likewise the

drones of Stock "B" are constitutionally the brothers of the worker bees in Stock "A". In breeding practice this means that if it is intended to use Colony "B" as a supplier of breeder drones, the following method must be used to determine their heritable value: a) Investigation of the physical characters of the drones of Colony "B". b) Evidence of performance of Colony "A" its sister and ancestor colonies. c) Investigation of the physical characters of Colony "A" workers. In this connection (i.e. use as a drone provider) the performance and appearance of Stock "B" are of no importance. So queen Y/81 can be apiary mated and though her workers may be sting happy and susceptable to Nosema, the quality of her drones will be unaffected. It is right to require that the virgin queens should come from stocks which have undergone full testing, but this is not necessary for stocks which are chosen as drone providers. Queens from the previous year, and, as just said, apiary mated, can be used without hesitation as drone providers, but on the other hand colonies from which these queens were derived, must have gone through the selection process. Only queen Z/83 will produce hybrid drones. "Normal" relationships are found in the male line by equating Drone = Hive Mother, except that there is a displacement of a generation, the actual grandmother of the drones is genetically speaking their mother, and her drone husbands are their fathers etc.

In the pedigree diagram of the official breeding form (e.g. on the Selection Report, Figure 23) the queen's mother and the mother of her mates are therefore both indicated as being at the same-generation level (for example 2a and 2b) whereas the drones are not shown as having their own generation. Herd book recording when several drone producing stocks are used: Drones from several sister queens are often brought to the mating station. There is therefore some uncertainty under which herd book number these drones should be recorded on the cards. The best solution is to record them under the grandmother's number as if she were their true mother. (For example if several daughters of queen X/80 (Figure 22) are employed as drone providers, the entry on the breeding card under the heading "Father Colony" Herd Book No. is X/80 D, where D = daughters). It is this queen (X/80) whose colony has shown the constitution inherent in these drones in respect of performance. If the breeding drones originate from more distantly related queens (1st or 2nd cousins) then only the number of the breeding line is shown on the card. Of course the actual number of each individual drone mother will be recorded in the herd book.

62 6. Breeding Programmes Anyone who operates his own mating station or offers an instrumental insemination service, is faced with the question: what should be the family relationships between virgin queens and mating drones in order to avoid too close inbreeding in the progeny? Experience shows that with bees, close inbreeding over a long period must be avoided, otherwise damage ensues which becomes apparent especially in a diminution of the rate of development. Closely inbred stocks produce too little brood and simply fail to prosper, so that in spite of the excellence of the rest of their genetic inheritance, performance deteriorates severely. The situation can best be made clear by referring back to the diagram in Figure 22. For the mating of queen Z/83 by line breeding, the following possibilities are usually open (that this queen has been derived from cross- mating is being disregarded for the purpose of this example). a) Mating with drones of the same stock (B) Relationship in accordance with family bond: Brother-Sister. Relationship according to genetic connection: Mother-Daughter. The degree of inbreeding with this mating is fairly high (25% increase per generation). b) Mating with drones of Stock "A". Family relationship : Uncle-Niece. Genetic relationship : Grandmother -Grand-daughter. It is true that this mating combination results in a quite low degree of inbreeding (about 12.5% in the first generation), but can rarely take place because queen Y/81 must be tested to enable her to be used as a breeder- mother, and it is doubtful if the old queen X/80 will still be alive. This is especially so when there is a two year test period between the generations. There is, however, a similar small degree of inbreeding if drones from the other daughters of queen X80 and even smaller if drones from the daughters of her sisters are used. This way out can be recommended for reducing the generation gap. By this means the very desirable possibility of impreg- nation by drones from several queens is realised at the same time. However, on practical grounds the opposite way is often followed; the old breeding mother is used again and the young queens are mated with drones of a later generation. c) Mating queen Y/83 with drones from Stock "B" (and other sister stocks). Family relationship : Aunt-Nephew. Genetic relationship : Siblings (Brother-Sister). There will always be sufficient numbers of daughters of the old queen to act as drone-providers, but by this mating system the increase in the degree of inbreeding is again as high as in Method A, about 25% in the first generation. It should be noted in this connection that matings of close relations at provincial mating stations even for a few generations apparently have no harmful effects on the bee. In particular the danger of gene impoverish-

ment has greatly diminished since it has become the rule to take several drone stocks to these Stations. Moreover, we know nowadays that even at a good Provincial Mating Station a quarter at least of the drones with which each queen is mated, are stranger drones; at the average Mating Station there will be even more. Harm from inbreeding is hardly to be reckoned with in such circumstances. However, it must be made clear that even at the Provincial Mating Stations one may carry out matings within very close relationships only for a very limited time. Working with precise mating control either with instrumental insemination (e.g. commercial insemination services) or at the Island Mating Stations then becomes very tricky. It is then very quickly realised that the honey bee is extremely susceptible to inbreeding, a fact that is closely related to the peculiar sex-determination of the honeybee and which will be briefly touched on in what follows.

The Sex Alleles Sex determination in all living creatures, leaving aside a few exceptions results form a uniform, very simple pattern. The heritable bases (the genes) are linked together in a number of bundles — the Chromosomes. The number of chromosomes is characteristic of each species, and each chromosome is present as one of a pair, one from the mother and one from the father. Every creature has thus two complete sets of chromosomes of a prescribed number, 2 x 23 in man, 2 x 4 in the drosophila and 2 x 16 in the bee. One chromosome pair, usually conspicuous both in size and shape, is unequal. One chromosome of this pair has a normal gene-stock, this is known as the X chromosome; the other known as the Y chromosome, has scarcely any genes. In these chromosomes the factors that determine sex are found. The union of XX produces females, XY produces males. The females produce eggs each of which contains an X chromosome, males produce spermatozoa half of which carry an X chromosome, and half carry a Y chromosome. From the fertilisation of the eggs individuals develop, about half of which carry XX and about half XY, which are respectively females and males. This is a very simple and reliable method for achieving equal proportions of the sexes. One of the few exceptions to what has been said is found in the honey bee. The proportions here are different, since drones are produced by parthenogenetic reproduction and possess only 16 nucleus coils in the chromosome set. Sex-determination according to the XX-XY system would be quite impossible, the system of sex determination in the bee (discovered by O. Mackensen and J. Woyke) is still, however, basically similar and just an exception to the general rule. In the bee it is not an entire chromosome, but a single gene, that deter- mines sex. There are about a dozen variants of this hereditary factor (gene) denoted by al, a2, a3 ...... a!2. This range of sex-determining genes is

called the Sex Alleles. There is a further difference in that the male has not a mixed inheritance, corresponding to X and Y; this occurs only in the female. If two different alleles come together (e.g. a3, a7) a female larva always results. If however, only one type is present (e.g. a3) then a male is produced. Since there is only one set of chromosomes in each unfertilised egg, and therefore only one sex allele, only a drone can be produced from it. In a very few cases from free mating and at a very much higher rate from inbreeding, it will happen that by pure chance two similar sex alleles come together (e.g. a3 and a3) in a fertilised egg. According to the general rule this union should produce drones, but the bees' scheme for living does not allow this; scarcely are the larvae hatched out of these eggs than the bees destroy them, perhaps because they have not the correct odour. Using one or two technical tricks (rearing artificially in an incubator, feeding on royal jelly) it is possible to obtain adult drones from such eggs. As theory would indicate they are very big, sterile drones. This loss of brood manifests itself to the beekeeper as "Pepperpot Brood" (Figure 24). Brother-Sister mating (drone and queen from the same stock) will result in an average loss of brood of 25%. By repeated back crossing with drones from the same source or by repeated brother-sister matings individual queens will be produced whose brood failure amounts to 50%. Part of this loss is made good by later egg laying in the comb, but

with such a high rate of wastage among the larvae normal development of the brood nest is no longer possible. The exact count of the holes in the brood is therefore of great importance for every pure line breeder and for every testing apiary. If only stocks which show very few holes among the brood are used for breeding, then queens will be chosen which have a greater number of sex alleles. (Figure 25). The consequence is to prevent these destructive manifesta- tions by which many a breeding line has been brought to ruin. Further details about distribution of the sex alleles and the technique of identification are to be found in the volume "Symposium on Selection and Mating Control, Lunz am See 1972" (obtainable from D.I.B. and O.I.B ). (German B.K.A. and Austrian B.K.A.) The counting of brood is very simple. At a time when the queen is laying well, place a brown comb of high quality in the middle of the brood nest. Three days later examine to see if a wide area has been laid in. 12 days after inserting the comb, count the unsealed cells using a template, with a rhomboidal opening which exposes 10x10 (=100) cells. (Figure 24). The mean will be calculated from a number of counts at different positions on the comb. If the count is delayed for a few more days, re-laying could produce a false impression of a well filled comb. In practical application the essential to be grasped is as follows. An individual queen in her gene-assembly has only two sex alleles, for example a3 and a7. She can therefore produce only two types of drones a3 and a7. But in her spermatheca there may be as many types of spermatozoa as

there were drones mated with her. If, therefore, not only this queen herself, but four of her daughters are taken as drone providers not just two but six different alleles are provided in the drones (4 from the mother's husband drones, 2 from the mother herself. If the drone providers are not sisters but cousins there may then be eight different types, almost as many as when free mating takes place. Hence not just the drones from one mother but drones from several mothers must be the rule! Only when the queen and all her drone-husbands possess different sex alleles will the brood nest be completely filled. Queen Drones al,a2. a3, a4, a5, a6, a7, a8. If by means of test crossings of inbred lines, the location of the individual sex alleles has been ascertained, it should be possible to bring about such combinations at will. (V. Maul 1972, Symposium, Lunz am See). In addition to the sex alleles there are — as in other animals — without doubt still more factors involved in inbreeding that have not yet been sufficiently investigated. One effect is common to all of them, that they weak- en the "life force" in colonies — sluggish build up, poor colony strength, poor summer yield, high winter losses, and all this in spite of careful

67 selection over generations for the best gene combinations. A sudden descent from the head of the Table to the bottom can take place from one generation to the next, simply because by a particular mating too many inbreeding factors have been accumulated in a queen. Recently, Page and Laidlaw in the U.S.A. and R. Moritz (Z.f. Tierzuchtung u. Zuchtungsbiologie 101, 1984) in West Germany have calculated with the aid of computer models, how large must be the number of breeder animals in order to remain as long as possible below the danger level for inbreeding. It was assumed that a dangerous reduction of pro - ductivity occurs with a degree of inbreeding of more than 25%, that is to say it is to be expected after a single generation of brother-sister mating. If queen rearing and mating control is practised (i.e. if every colony is not free to raise and mate its own queen) a certain degree of inbreeding is inevitable, since all the queens in the apiary are derived from a small number of breeder mothers. The extent of increase of inbreeding per generation depends on the number of breeder animals (queen mothers + drone mothers) employed. It was calculated that, if eight breeder animals per generation were used in line-breeding, 10 generations would pass before this boundary would be reached (Figures 26 and 27). If the method set out below were followed that would be about 15 years (or 20 years if 10 breeding animals were used). With the interjection of Line-Combination however, as Bienenfeld was able to demonstrate on the basis of results collected in West Germany, the degree of inbreeding can remain permanently at a very low level. But on the other hand, he was able to deduce from the same data that a decline in yield ensues from a degree of inbreeding substantially less than 25%. But in a similar way the harm can be remedied in a single generation or at most two, by suitable mating. Why it often takes two generations to get rid of the harmful effects of close inbreeding, is quite simply explained in that two generations similarly play a part in the performance of a colony of bees, namely the queen and her daughters, the worker bees. If an inbred virgin queen is mated with drones of another strain her daughters will of course display hybrid vigour, but the queen herself remains weakened by inbreeding and so will be unable to build up a colony to full strength, because of the reduction of her egg laying ability and the defective constitution of the eggs she lays. V. Maul has made a very impressive demonstra- tion of this by crossing two inbred lines which nonetheless possessed the appropriate combination of sex alleles. (Apidologle 1982/1). Not until this queen's daughters were suitably mated were they able to display in the workers they produced, and in themselves, complete vital energy. The breeding plan for every line-breeding programme is of critical importance, therefore. Anyone who sends his queens to outside mating stations need have little fear of inbreeding, since the drones are not of the same strain. On the other hand he should take care to remain within the same kinship group (or breeding strain) since otherwise the outcome can scarcely be foreseen.

Hybrid breeding within the race (and therefore crossing of different strains) often provides outstanding results; but many times results are dis- appointing. Because of the unknown, unforeseeable results, only those who have a wide range of selection possibilities at their disposal, namely experienced breeders at an Institute, or a Testing Station, or a large private apiary, should venture into this field.

A Reliable Programme for Line-breeding Groups of breeders or individual breeders who wish to maintain and increase breeding Lines either at their own mating station or by means of instrumental insemination will have to give serious consideration to the manner in which they must organise their breeding operations so that they will be able to operate without diminution in vitality and productive capacity of the colonies in the long term. There must be a minimum of at least 60 colonies available as a base, since otherwise sooner or later the ravages of inbreeding will make their appearance. The following breeding plan may be regarded as a pattern to be modified as necessary to suit prevailing conditions. The starting point is a group of queens from a reliable breeder whose breeding stock has proved itself up to now to be suitable for the existing forage conditions and system of management. A start is never made with a single queen; only with a group of five or still better, ten queens, is there adequate scope for selection. If possible they should not be sister queens.

The basic idea of the breeding plan is that three more or less independent Lines will be developed from the starter group. Year by year drones from each of these Lines in turn are taken to the mating station. By mating with queens from their own Line, this Line will be preserved (Pure-Line breeding). Mating with queens from the other Lines produces Line Combination. The breeding plan represented in Figure 28 proceeds as follows:- The first year after the purchase of the queens is employed in testing in accordance with the usual principles (Page 53) and in rearing small test-batches from several mothers. On the basis of this performance test three breeder-mothers A, B and C are chosen together with other queens or, as appropriate, their daughters as Drone-mothers.

First breeding year; Rearing of large batches from each of the three queens A, B and C mating with drones from several of the 10 original queens or from their daughters. These three batches of sister-queens are the 1st generation of the Lines A,B and C. Second Breeding year; Drones from Line C Queens 1. from Line C (Pure-line breeding)

70 2. from Lines A and B (Line combination). Third breeding year; Drones from Line A Queens 1. from Line A (Pure-line breeding) 2. from Lines B and C (Line combination). Fourth breeding year; Drones from Line B Queens 1. from Line B (Pure-line breeding) 2. from Lines A and C (Line combination). In order to shorten the intervals between the occasions of Pure-line breeding, small batches of Line-mated queens can be inserted into the programme taking each Line in turn by using instrumental insemination or by sending them to a second mating station. As more than thirty years experience has shown, while continually selecting for productivity, vigorous development and quality of the brood nest, continuous breeding can be practised for a very long time by this system without having to contend with any harmful increase in the degree of inbreeding. The consistent use of an adequate number of breeder mothers (as shown in Figure 28) is important; at least three mothers per Line for the queens and six for the drones. A programme of this kind requires some outlay, especially in colonies. It will be carried out best by a sharing of work, perhaps by a group of breeders, but at least as the joint work of two. Since the introduction of artificial insemination services, which allows the use of a variety of drone groups in close proximity to each other, the possibilities are greatly increased. The breeding plan outlined here which brings about mating between demonstrably related partners is designated "Pure-line Breeding". Since the testing of queens is mainly carried out with groups of sisters of the same age, this kind of breeding is very common. This is the quickest way to a uniformly good stock of breeding material. It has been shown above how in spite of mating between relatives, the harmful effects of inbreeding can be avoided by the use of a number of separately developed lines.

The aim of Pure-line Breeding is twofold:- Guarantee of rigorous selection within an extended family of known descent (= within a closed population) while at the same time maintaining the lowest possible degree of inbreeding. Line-breeding is therefore funda- mentally different from other breeding systems. Above all it avoids "jump- ing about" from one source of breeding material to another, and the continual crossing-in of outside strains or races which is all too common in the beekeeping industry. Both may indeed produce ephemeral peak harvests, but long term success can never be achieved by this means. In the long term inbreeding can be kept low by the division of a breeding strain into several breeding Lines (Bienenfeld 1987). This system is primarily suited to practical conditions: A breeder or a breeding group, which for example maintains three Lines of a strain, is thereby constrained to make use of many breeding animals on a regular basis. Practical experi-

71 ence has shown that this method can be worked for a very long time in a closed system without deterioration of brood production. In all the cases I have known where just as a matter of convenience the Lines have been combined after a time into a single group, signs of inbreeding have appeared sooner rather than later, and it has been necessary to bring new breeding material into the apiary. If however, several Lines are available, possible inbreeding effects can quickly be corrected by crossing between the Lines; and by subsequent back crossing, the original Line can be restored in its essentials (breeding in "half-isolated groups" according to Professor Pirchner, Weihenstephan). In practice it has proved satisfactory to classify the Line as a "Line", if there is at least 50% of this Line in the queens. The Lines of the Carnica strain "Troiseck" have been bred at Lunz am See since 1948, that is for nearly 40 years, according to this system with increasing vitality and productivity. During this period these Lines have become so widely distributed that there are numerous sub-Lines providing possibilities for even more combinations within the Line, and so providing a still broader basis to the work. Up to now, similar results could not be obtained by any other system. The breeding scheme put forward here has thus fulfilled in practice the expectations placed in it. Along with honey production, the egg laying performance of the queen together with the compactness of the brood nest, must be well to the fore among the qualities for selection. By attention to this, selection continuously works against inbreeding. If damage from inbreeding appears in a previously good strain (e.g. unsatisfactory build up of the colony) a simple remedy is readily available by using drones of a different Line or Sub-line of the same strain for mating. But as was indicated previously, clear improvement will often only appear in the second generation. By suitable combinations of breeding Lines, outstanding foraging ability can very often be united with great vitality and lively build up. This breeding method is recommended with special emphasis. When animals are mated which belong to the same strain, without being very closely related (as for example, when an outside mating station of the same race is used), it is usually called Pure Strain Breeding. If the pair do not belong to the same strain, but only to the same race, it is called Pure Race Breeding. To make the matter clear, the most usual terms which refer to mating are defined as follows;-

Breeding Line Groups of queens of similar geographical origin, which have an authenti- cated relationship with each other. They can also be called an Extended Family or a Kin-group, or a Population. Well known breeding lines were, and are, for example;- Sklenar 47 and Troiseck - Lunz 1075

Breeding Strain Queens of particular origin and descent, but not closely related to each other. Among the Carniolans widely distributed strains are;-

72 Troiseck from Styria, Sklenar from Lower Austria and Bukovsek from Stovenia. Races Bees of defined geographical origin which are characterised by the same body — and behavioural-characters. Races can be distinguished from each other by these characters, which is not the case with Breeding Strains or Breeding Lines. Line-breeding is the mating of animals which mainly belong to the same breeding Line, as set out in Figure 28. Since it is scarcely possible to keep a line completely closed for long, but combinations with other lines of the same strain have to be made from time to time, there needs to be agreement as to when it can rightly be called Line-breeding and when not. It has proved advisable to use the term Line-breeding for matings in which the proportion of the line in question amounts to at least 50%. If, following a line combination (such as BxN Figure 28), there is a back-cross in the next generation with pure, or predominantly pure B drones, then the principle of the Pure Line is preserved.

Racial Hybrids result from matings between animals of different races Work with "crosses" or "Hybrids" has had great successes in many areas of plant — and animal — breeding. According to previous experience similar results should, in principle, be obtainable with bees. Consequently, the method of crossbreeding has been recommended by many as the most promising way to success. These recommendations are both shortsighted and ill considered; for conditions vary, very considerably, from district to district — especially as regards bee breeding. Cross breeding (see p. 98) will undoubtedly become increasingly important in many regions; but the assumptions as regards Central Europe are unsound on the following grounds. To establish hybrids, two lines or strains will first have to be selected, which together will produce a good combination for the district in question, since not all crossings are certainly superior, nor even of equal value. These strains will, therefore, have to be made as uniform as possible by selective breeding, so the desired results of crossing fails. Furthermore, much preparatory work is needed, which can generally be carried out only in a research station. According to an abundance of previous experience, the hybridisation of races without previous selection of the partners for the cross-mating very often leads to disappointment. (K. Weiss and colleagues: "Beekeepers' Friend" 1981 pp 75-90). Furthermore, at best the hybrids can only be raised by the greater breeding businesses, since for the purpose, two foreign races of bees, and a reliable mating station are needed. There should be no breeding from hybrids. This requires that a new queen will have to be bought by the beekeeper for every stock at least every second year. This outlay would be paid for only if a long continued

Figure 29. Later generations from a "Blender" regularly show a decline in productivity (explanation in text). increase in yield of 30% amounting to at least 15Kg per hive per year, could be relied on. It is one thing to work in a country where these conditions can be confidently expected. It would be different in our climate. What is more, with the introduction of Racial Hybrid breeding, only a restricted circle of progressive professional and semi-professional beekeepers would be interested in the business of hybrid-breeding — and in all probability not for long. The great majority of small beekeepers would not be interested. The final result would be a total collapse of all attempts at improved breeding, and an uncontrolled mixture of sting-happy cross- breds which would make beekeeping impossible in many areas. The pre- conditions for the use of racial hybrids are good, wholly reliable nectar flows, and a beekeeping structure in which business enterprises predomi- nate — not our small apiaries, and uncertain, modest nectar flows. The introduction of the Carniolan bee into both West and East Germany has — along with other measures — resulted in a considerable increase in the average yield during recent decades. For many years before that, unco- ordinated trials and all possible kinds of imports (a few gamblers still do it today!) had been carried on. But the conviction has finally prevailed that national bee breeding can only thrive with a uniform, well adapted, peace- able bee. This understanding arrived in the nick of time, for in future the public will refuse to accept a bee with the aggressive nature of the old local bee. All breeding procedures are based on the assumption that the Father is selected equally with the Mother. If the extraordinarily strong influence of the father bee is not taken into account , then nothing more can be said

about breeding, for then the breeding qualities would change in a completely uncontrolled manner, from generation to generation. Nonetheless, apiary mating has its important place in methodical queen rearing; the easily and cheaply reared daughter of a tested pure bred mother can give outstanding performance as "working" queens. It is important that there should be no further breeding from them and that the pure bred line remains as the source of breeding material. These "working" queens are mainly cross-mated, since the mates come from a different strain, but they are very cheap, and can be obtained in almost unlimited numbers. If the mother of these queens comes from a tested strain, their performance will be good, and, since they have mated outside the strain, their colonies will be very vigorous. But where there is a strong input of "black" blood from the local strain, the "working" stocks may be restless and not always quite gentle. But the stronger the Carniolan influence in the district, so much more gentle will the first generation

75 crosses be. The ideal is the rearing of "working" queens in a pure-breeding area. Beekeepers who prize stocks capable of good performance and reasonably good tempered, but who are either unwilling or unable to under- take the trouble of selective breeding would be well advised to buy a good supply of good queens, of pure strain and purely mated. From the best of these, pure-bred mothers can be obtained when needed. Mating takes place at the apiary. Subsequent breeding must only be from pure-mated queens, which form the breeding strain. Most of the colonies are headed by locally mated "working" queens. Any beekeeper who does not wish to rear queens himself should obtain "working" queens from a reputable breeder just as it has long been the practice among breeders of poultry. It cannot be too strongly emphasised that there should be no requeening from locally mated queens and that no swarm cells should be allowed. If two or three neighbouring beekeepers can be persuaded to adopt the same procedure a small "pure breeding zone" may be formed very quickly. The only proper procedure for Central European conditions is pure breeding within a race. "Pure breeding" certainly does not mean "inbreeding"; with systematically organised line breeding there is no longer any danger of debilitation from inbreeding. Moreover, a geographical race like the Carniolan embraces so great a variety of individual types that there is ample scope for combination breeding within the race. For areas with mongrelised local bees the breeding aim is "Displace- ment breeding by means of Carniolan Lines". Eventually we should reach the point where even apiary mated queens will retain in their colonies the essential characters of their own lines, and the drones of the district will predominantly transmit the attributes of the same race. Only then will the efforts of the breeder be of benefit to the broad generality of beekeepers. This goal cannot be reached by pure bred queens alone, but only through their apiary mated daughters. Bee breeding is therefore carried out in two stages:- 1. Provision of pure-mated queens from a tested line as breeder-mothers. 2. Rearing of apiary mated daughters of these queens as "working" queens. Each group has a separate purpose; The employment of a genuine pure mated queen just for honey production and not for breeding is just as wrong and uneconomic as breeding from a top class "working" queen. The old rule that one should never breed from hybrid and production queens (i.e. from "Blenders") no matter how good their performances and qualities is borne out again and again. One example from our own experience must suffice. (Figure 29) Queen 'P' was a pure Sklenar Queen with good performance. The colony of an apiary mated daughter (HI) was for three years in succession both in development and performance, far ahead of all the rest in a large apiary. Since there was no serious objection to the colony as regards physical characters, the queen was used as a breeder. All the subsequent matings took place at the mating station. The next generation (H2) was generally

above the average for the apiary, but not one colony reached anywhere near the level of HI. In the third generation, performance had generally fallen well below the apiary average; among 16 annual production reports there was not a single one which would have suggested that the colony was worth breeding from. On the other hand, that the productivity of one and the same population can be maintained at a high level over many generations and even raised still higher by properly managed Line-breeding, is shown by the "Troiseck" strain. Lines ("07" , "1012", "1075", "Wintersbach") bred according to this principle at the breeding station at Lunz am See since 1948. The most convincing proof (on account of the large number) of the positive results of long term breeding can be inferred from a summary and analyses of the records of the Testing Apiaries for Carnica bees in West Germany and Austria for the years 1961 - 1983, by K. Bienenfeld (Figure 30); 6187 Annual Returns of colonies were included. The five-year averages of honey yield show - with fluctuations related to nectar flow — a

77 continuous increase in yields from 18.7 kg in the first period to 25.4 kg in the last, (an increase of 6.7 kg). The increase in yield of 0.25 kg per year for about 25 years could indeed result in part from improved management, while forage supply during this period has certainly not improved but on the contrary, as a result of agro- technical measures it has grown worse in most districts. The main part of this increase in productivity, which lies well within the framework of results in other areas of animal breeding, can consequently be ascribed to a consistent selection procedure. Wintersbach strain of the Troiseck line has been a peak performer for more than 20 years without ever receiving new "blood" (i.e. new gene combinations) from outside (only combinations with related lines). As expected the yield fluctuated about the 100% mark, the results are uniform. Occasionally (1962, 1972) inbreeding-depression occurs. But this is corrected by back-cross to other breeding colonies, or by Line combination. The latter often produces "Peak output as planned", but the Breeding Centre has to incur expenditure in order to obtain the outside Line.

78 Chapter III Control of Mating In practice, controlled natural mating can only be achieved by taking the virgin queens and selected drones to places so isolated that they can hardly be reached by stranger drones (Queen Mating Stations). Alternatively such an enormous preponderance of the preferred strain of drones is produced, that outsiders have only a remote chance of mating. (Mating Apiaries and Pure-Breed Districts). Because of their inherent insecurity these last named systems are not accepted as being "Pure breeding" but under certain conditions, which are explained more fully below, they can result in a very uniform even tempered bee and can make a substantial contribution to improving performance in the locality.

1. The Mating Apiary Controlled mating can be achieved even in the apiary under the following conditions;- a) The distances from adjacent apiaries should be not less than 2 to 2.5Km, according to the formation of the land in the locality. b) A large number of colonies in the apiary (40 to 50). c) Requeening all the colonies in the apiary with queens of a prescribed Line, possibly the daughters of a single queen. d) Systematic breeding of drones in these colonies. The situation is all the better, the smaller the number of stocks in an area or radius 2-4 Km surrounding the apiary. Success can be seen wherever one beekeeper's bees dominate the whole district and neighbouring apiaries are stocked with the same breeding material. It depends on the fact that drones of the desired constitution can be quickly produced in great numbers — simply by requeening with tested (selected) queens in great numbers. These are allowed to mate in the apiary and are therefore production queens. The great number of drones is obtained by placing a drawn out drone comb in every stock with good performance. This remains all summer and no drone brood is removed. We know nowadays that this procedure, while continuously maintaining a great army of drones, does not in the least reduce the honey yield. The neighbouring beekeepers, having been supplied with the same material, are then advised to adopt the same system. By this means a small pure race area is established, which with continuous effort over the

79 years is extended ever further. I can cite a number of examples of beekeepers in sites that are far from being isolated, who have made such progress by these methods that the results of testing of their apiary matings are scarcely distinguishable from those of the Island Stations, as regards the physical characters of the bees. Many of our conventional District Mating Stations (with a Bee-free vicinity frequently of only 2-3 Km radius) are after all very much the same, but with this difference; instead of an apiary with 20-30 stocks, there will only be 6-10. But in support of the Mating Station, there is usually either an entire Association, or a group of breeders, for whom no trouble is too great in order to create a broad pure race cordon. In districts which have a high bee population, a Mating Apiary makes no sense, unless it is totally protected by such a pure-bred belt.

2. The Pure Breeding Area. By extending the pure breeding cordon described above, a Pure Breeding Area can be created. In many districts where the bee population is dense, the creation of a Pure Breeding Area, is the only means by which matings with drones of the same strain can be secured. This is the quickest way to get preferred bees established permanently. It is not possible to give in detail the specifications for creating a Pure Breeding Area. Much depends on the site of the apiary the number of stocks, etc. The minimum acceptable requirements for a Pure Breeding area is a distance of 8-10 Km (better still 15 Km) and an establishment of 300 colonies. Creation of such an Area depends less on the inclusion of every apiary (in which case most such projects would fail!) than on the determination with which it is undertaken. Over a period of years queens of pure descent must be continuously poured in. To start with, there must be no requeening from locally mated queens, but only from pure bred mothers bought in for the purpose. In the course of a few bee-generations, those apiaries inside the Area which could not be requeened, will have changed quite spontaneously. Outside the area, a mixed population belt develops, in which the inheritance of the pure race has a substantial share. Experience has already shown that in a Pure Breeding Area with 500 stocks a very satisfactory degree of pure mating is achieved - apparently better than in the ordinary mating stations. When the Pure Breeding Area has reached this condition, and for several years has exerted an extending influence on the surrounding apiaries, queens can be offered for mating in the centre of the district. Breeding selection in the proper sense is certainly more difficult and slower, but once the chosen line has been established, it can be maintained unaltered over a long period. In a Pure Breeding Area, selection for breeding at first only occurs in the mother-line. In the next generation the father-line is also affected by it. If

80 drones from the selected mothers are to be acceptable, they must therefore be bred only from the most uniform and best stocks. Drones from hybrid stocks with poor performance can be eliminated by fitting a drone excluder to the flight hole of these colonies during the period of mating flights. By using a grid-width of 5.2mm (obtainable from Firma Heuner, 8532, Bad Windsheim) the foragers can fly in and out quite unhindered. The screen is so placed in relation to the flight hole, that it slopes downwards and the drones do not push so hard against it and there is no congestion at the flight hole itself. (Figure 34, p. 89). Moreover, drone breeding in these stocks is continually hindered by cutting out the combs, while drone breeding is encouraged in the good stocks. Furthermore, daughters of newly tested pure bred mothers should be brought in to the Pure-Breeding Area again and again. In the Pure Breeding Area all the requirements for the raising of production queens are provided. This type of queen, which forms the backbone of a successful honey industry, and should therefore be used more extensively than formerly, can be characterised as follows;- 1. Derived from a pure-bred mother of a tested breeding line, and therefore with a constitution guaranteed for good performance and gentleness. 2. Mating predominantly with racially similar drones, resulting in exceptional uniformity. 3. Mating takes place in districts where the climate is favourable so that results are good in number and quality. (Complete mating). 4. Little expense is involved, and so good value for money. 5. No danger of sudden decline in performance because of inbreeding.

3. The Mating Station It should never be forgotten that at any Inland Mating Station, only the conditions for pure mating can be provided in accordance with the knowledge and resources available. What actually takes place there is never known in advance. This can only be inferred later from the behavioural and physical characters of the progeny. "Mating Station Mated" does not necessarily mean "Pure Mated"; not by a long chalk.

Sites In the last issue (1986) of "Breeding Regulations of the German Beekeepers Association (DIB) two types of inland Mating Station were differentiated:

1. Line Mating Stations. "The distance from the nearest apiaries with drones which are undesirable for the breeding programme must be at least 6 (better 7-10) Km. This belt is designated the Guard Ring of the Station. There must be a firm guarantee that all colonies established in the Guard Ring are provided with queens derived from selected Breeder-mothers of the Breeding-Line of the Station".

81 In contrast with the previous rules, three points in particular are new: There is no mention of a "Safe site". This indicates that isolation depends in fact primarily on distance, and the number of colonies, and less on topography. The minimum distance to the nearest "stranger" drones was increased from 3-4 Km to 6-10 Km. This change is the result of new investigations into the distance flown by drones, and numerous pieces of research into mating, carried out with "Cordovan" mutants, which have revealed far greater distances for mating than was previously accepted. Instead of speaking of "Bee free zones", it is now possible to talk of "distance from stranger drones", for in fact it is the drones, and not the bees, which determine the result. It is not therefore necessary for an area of at least 6 Km radius round the Station to be free from bees, but they must be just as well managed and provided with the same kind of queens as the colonies at the Mating Station. Only a few Stations in the mountains and a very few in the lowlands will comply with the required conditions, that is to say, in places where the beekeepers of the area are fitted completely into the breeding programme. Every Line Mating Station should sooner or later be tested for its reliability using the Cordovan Test (p.93). The necessary instructions can be obtained from the nearest Bee Institute. In the choice of a site and in the

82 management of the Station, careful consideration must be paid to the increasing information about flight distances and direction of flight of queens and drones. In the crafting of the regulations governing Mating Stations, a good deal of freedom is quite deliberately allowed for the assessment of each situation individually. The absence of rigid specifications and better understanding of the procedures will lead to more reliable results. Island Stations are obviously Line Mating Stations, and in quality greatly surpass the Inland Stations. On the basis of present experience, a distance of 3km over water from the mainland is sufficient for isolation.

83 If there are mating stations for different Lines on one island, the regulations for a Line-Mating Station on the mainland are applicable. (The distance between these stations to be at least 6km). What was said about the choice of site and setting up of Inland Stations applies here, but special attention must be given to protection from wind, and heat-insulation. Nowadays, it is regarded as self evident that there should be a sufficient number of well nourished drone stocks (never less than 4) for the wear and tear on the drones is naturally greater on a North Sea Island than on the mainland. Instrumental insemination of queens is included in the category of Line Breeding. The technicalities of preparation of queens and drones are dis- cussed in the brochure "Instrumental Insemination of Queen Bees" 2nd Edition 1973, Apimondia Publishers, Bucharest (Figure 32). Instrumental insemination of queen bees has acquired growing importance for practical beekeeping in recent years, especially since it has been shown that inseminated queens are not inferior in performance to naturally mated queens. (F. Ruttner, "Beekeepers Friend" 1976/ 11 pp 351-3). The results of inbreeding of the honeybee only became apparent to a marked extent when a genuine control of mating became possible with the setting up of Island Mating Stations and the perfection of the techniques of Instrumental Insemination. Previously, Nature had taken care that faulty breeding plans were rectified so that no serious inbreeding could take place. With the employment of precise mating procedures it is more than ever important, that sound, well thought out breeding plans are followed.

2. Race Mating Stations. "The foregoing specifications (i.e. for line-mating stations) may be modified so long as the racial standards for mating are guaranteed". The Breeding Selection Superintendent and the District Chairman for Breeding have to make the decision whether an establishment is to be graded as a "Race Mating Station" or a "Pure Breeding District", depending on the conditions obtaining there. For a Race Mating Station no minimum distance from stranger-drones is laid down, the participation in the mating of racially similar drones from the vicinity being accepted, but it is of course assumed that the Pure Breeding zone round the Mating Station is subject to continuous supervision and selection with regard to performance. Otherwise, it should not even be called a "Mating Station" — therein lies the distinction from a Pure Breeding Area. Experience has shown that Mating Stations operated for decades by capable breeders, often produce very good results as regards both the quality and the racial purity of the progeny, although the isolation of these stations is by no means complete. The bee-colonies in the neighbourhood have however in the course of time simply been completely requeened so as to conform with the breeding aim of the mating station

84 so that racially similar matings have been achieved almost exclusively. That is why the work of these Mating Stations with their frequently very large despatches is entirely in line with the current breeding objective: what is most important at the present time is the rapid distribution of the gentle Carniolan bee to all places where remnants of the excitable local bee still persist. The mating results from the Stations must obviously be tested every year, at the very least by means of samples at a Selection Station, or by a few examinations of physical characters. The discoveries during recent decades about the biology of the mating of bees, and consequently on the performance of mating stations, have certainly destroyed many illusions besides laying the foundations for tidier, better conceived operations. For Line Breeding, for which precise control of the paternal side is essential, there are available Instrumental Insemination, Island Mating Stations and a few Inland Stations. The bulk the of the existing Inland Stations provide the conditions for the broader work of supplying selected, pure breeding material to the Beekeeping industry as a whole.

Establishment of a Queen Mating Station The site for a mating station should be chosen, wherever practicable, so that there is an approach road right up to the stocks. The mating nuclei should be set up in a "well sign posted" area (only single trees or bushes) and not too closely spaced (minimum distance between nuclei about 3m, or 5m if there are no landmarks), while uniform straight lines should be avoided. Queens perish mainly by drifting. A very well attested help to direction finding is to vary the height of the boxes above the ground, and the direction of the flight hole. For many decades now we have proved the value of stakes made from steel T-sections for securing the boxes. (Figure 33). They seem to last for ever, and can be driven in in a moment (even into coarse road metal) and can be removed when necessary. In loose sand they do not provide sufficient support. If there are ants at the site, the use of fruit tree grease is recommended. The roofs of the insulation boxes should always be painted with oil paint. Boxes with the same colour combinations should not be placed close together. There should also be colour markings above the flight holes. (III. 1.) Because of our changeable weather — especially in June when it can be quite cold — siting in hill sun is recommended, except perhaps in very sheltered and very sunny positions. By setting up in the shade crucial hours of sunshine can be lost. Fear that the little colony will abscond is unfounded so long as the mating boxes are correctly filled, and properly insulated covers are used. (III. 3.). The outercases must be lined with 12- 15mm toughened polystyrene on all sides and beneath the roof, so that they are adequately protected against both cold and heat. Old outer covers which are not big enough to allow this are better discarded.

The Mating Nuclei In Central Europe the size of the mating colony has been progressively reduced till suddenly we ended up with Peschetz's miniature E.W.K. (Ein Wabe Kastchen — Single Comb Box). Surely this has gone too far; generally speaking it is only for the more important breeder who produces on the larger scale for the market. The units which one raises for one's own use, are certainly not lost, and anyone who objects that he is short of bee material, simply shows that his management methods need to be overhauled. In Austria and in Germany during recent years there has been a reverse tendency towards larger mating nuclei, and a return to multi-comb boxes. These nuclei stand up better to periods of bad weather in the Mountain Stations (in particular, they stay healthy!) and have enough bees to form the kernel of a new stock. They are also well adapted for travelling, and if one considers the great reduction in the previously quite enormous losses during introduction this method is certainly more economical than using the smallest units. There is a whole series of multiple comb boxes either free standing or suitable for the usual containers. In recent years the Kirchhainer Mating Box has enjoyed an increasing popularity (Figure 13 and 14). As with the

86 old Swiss Mating Box no frames are used, simply three wooden bars fitted with starters. Examinations and the removal of the queen are thereby made very much easier. For transport over long distances care is of course required. The boxes are made of dense polystyrene and are therefore very warm, light and not expensive. The so called "Kernel-nuclei" have proved most effective as larger units, specially recommended for the beekeeper who breeds largely for his own requirements, but also for the breeder who prefers to deliver his queens in nuclei. They are made up from one comb of sealed brood, well covered with bees, one comb of food, and one frame of foundation and a ripe queen cell. These nuclei develop rapidly and with a little reinforcement can by Autumn provide a reserve stock fit for wintering without the queen ever having had to change to a different colony. The multicomb box runs into no difficulties at Mating Stations which are used by restricted membership breeding groups or by a single breeder. Generally speaking however the big Mating Stations accessible to all and sundry will have to remain, limited to the use of EWK's as formerly. The filling of the mating boxes seems to me even more important than the type of box used. Every Station Warden has had to report experience of two breeders who bring queens to the Mating Station on the same day; one has excellent results, while the other loses more than half his queens. This is certainly no accident. We know how very much the bees in the mating box influence the mating flight. Therefore strict attention must be paid above all to the proper age composition of the nucleus. Young bees in plenty! Losses caused by queens going astray on their homeward flight can be reduced by appropriate siting and marking of the boxes. Generally speaking, absconding of the mating stock should never occur; it is always the result of a technical fault — poor insulation of the outer cover; too many old bees in the nucleus. The queens should be marked if possible before the mating flight. That marked queens are more easily lost is a fairy tale; losses during the mating flight are very rarely attributable to birds. They occur because the queen cannot return because of cool or windy weather, or from failure to separate from a drone; or because on her return to her own or a stranger nucleus, she is stung to death by the bees. More often a queen that has gone astray survives, but she can only be recognised if she bears a numbered disc. A decision has often to be made how long the queens are to be left at the Station. The Station Director has to wait till the last queen is mated (or lost!) before he can return the whole batch. If, during the course of the 10- 14 days that the nuclei were at the Station, two or three days of good mating weather were recorded, it would not make sense to wait for one or two stragglers. They will not be worth much, even if in the end they are mated and it is better to abandon them straight away, than to harm or endanger the rest of the queens. Experience also shows, that with queens in small mating boxes which have not enough space for egg laying, very often the development of the

87 ovaries regresses to such an extent that they become liable to fly again, and further matings even take place. For this reason also, queens would better be brought to the Mating Stations in larger units.

The Drone Colonies 100 queens require approximately 1000 drones for mating. During the mating flights there must be available at the station 6 - 10 times this number in order to ensure quick matings near the sites of the nuclei. But since one stock can at the most support no more than 2000 drones at a given time (of which perhaps only half are sexually mature) several drone colonies must of necessity be set up. It has become evident at the North German Island Stations that there is a direct relationship between the number of drone colonies and the number of successful matings, the fewer the number of drones, the lower the proportion of mated queens, and the poorer their quality (Tiesler; Apimondia Symposium Lunz 1972). The number of drone stocks required at a mating station must be decided separately in each case, depending on the prevailing circumstances; the site of the Mating Station, the number of stocks of bees at the boundary of the Guard Ring, the number of queens to be served there. The numbers set out below are therefore to be taken only as approximations. The lower numbers in each case apply only to the most favourable sites, with the nearest apiary more than 6Km away, and with few stocks of bees in the district. For 50 Queens brought in at the same time (25 cover boxes in use) 4-6 Drone Stocks For 100 Queens brought in at the same time (50 cover boxes) 6-8 Drone Stocks For 250 Queens brought in at the same time (125 cover boxes) at least 8 Drone Stocks For 500 Queens brought in at the same time (250 cover boxes) at least 12 Drone Stocks At our most common type of Mating Station, (with only a 4Km drone free radius and many stocks of bees at the edge of the area) at least 6 stocks of drone producers are needed for 50-100 queens installed at the same time. But every additional one is an advantage! This follows, not only from the essential requirements for pure mating, but also from the recent calculations of the number of breeding animals required for keeping down the level of inbreeding. The Breeding Regulations (1986) require for Inland Mating Stations a minimum of six to eight drone producing stocks. These stocks must all be selected. It goes without saying that at a Line Mating Station all the drone producing stocks must be derived from the same Line. Experience shows that it is often difficult to provide the required number of selected drone stocks for the Mating Station. These colonies are also needed at the apiary for breeding and they should in addition be demon- strating their production capabilities in comparison with the other stocks. In this case a remedy can be provided by setting up drone-nursing stocks

from sieved bees and brood combs without drone brood. They must have at least 8 frames thoroughly covered with bees, and be provided with a queen cell ready to emerge and a comb of sealed drone-brood. The drones should be derived from several breeding mothers. As everyone knows, a queen in a vigorous stock can be induced to go on laying continuously in drone comb. The number of drones that a colony can rear is limited, however, so that when the larvae have hatched out, the combs must be removed and distributed to nursing colonies for rearing, being placed there between brood combs in upper chambers. As was mentioned on page 79 the number of drones can be greatly increased by the requeening of the stocks on the outskirts of the Mating Station district with queens of the chosen breeding Line. But these must all be selected drones, and every year before the start of the breeding season, the presence of the original queen must be verified (clipped wings, numbered disc). If the beekeeper concerned is prepared to co-operate this provides the most sensible way of increasing the number of drones, and at the same time the Station safe-zone is extended still further. As long as the drone producing stocks are still at the Apiary the entry of any foreign drones, and the drifting of one's own drones must be diligently prevented. This is done by setting up the stocks at least 100 metres from the nearest colonies of bees or — even better — by fitting a drone trap (Figure 34) with an excluder 5.2 mm mesh width sloping downwards and inwards. This closure does not hinder worker bee flight in the least, and the drones do not struggle much on the screen because their view of the open sky is cut off. (In addition, such excluders are also very well suited for stocks which happen to be in critical proximity to a Queen Mating Station). Excluders with a mesh width of 5.22 mm are obtainable from Firma Konrad Neuner, 8532 Bad Windsheim Bavaria. These drone-devices have the further benefit that flying activity can be readily observed in them and they are also convenient for obtaining drones for instrumental insemination. But as soon as the maturing drones are ready to fly and press against the screen on warm days, as a result of the

89 large numbers a great press forms and if it continues will do injury to the drones trying to escape At this juncture, the colonies chosen to be drone producers should be moved immediately to the Mating Station and the drone traps taken off. But if this is not possible, the drones' urge for freedom can be better curbed by restraining them in the colony behind an excluder; in the honey chamber in side opening hives and above an excluder placed between the floor board and the first box in storey hives. Of course, the crowd of drones cannot be kept here indefinitely. In stocks at the outskirts of the Mating Station area, from which drone flights should be prevented, the number of drones can be kept small by regularly cutting out the drone comb. Bees and brood for the drone-rearing colonies come from reserve stocks which were prepared for wintering the previous Autumn specifically with this in mind. At the Breeder's apiary there ought to be a substantial number of nuclei which have been made up in good time; only when the bees are available in ample quantities can the work proceed without check and in a really satisfactory manner. It is recommended that a good drone comb be placed in the middle of the winter-clustering space of the colonies intended for drone-production before autumn feeding begins. The queen will probably lay in the drone comb very early in the season, so there will be time enough to carry out the examination of physical characters. More colonies than will be strictly necessary should be prepared in this way, to compensate for losses (and possibly unsatisfactory character results). Drone rearing can be even further advanced by stimulative feeding with candy fortified with protein (but on no account before the sallows bloom!) (for further details see the time table p. 44). If still more drone-colonies have to be formed from one particularly good breeder-stock the procedure is as follows:- If larvae are present in the drone comb, it should be removed and placed between brood combs in the honey chamber of a very strong nursing stock. If necessary this stock should have been strengthened in preparation. The breeder colony receives a new drone comb without delay.

Setting up Drone-nursing stocks Two days before the emergence-day of a drone comb, a brood-nucleus is prepared in the usual way with the following composition and order;- 1 comb honey and pollen. 2 combs emerging brood, well covered with bees. 1 comb sealed drone brood together with bees. 2 sealed brood combs, well covered with bees. 1 comb pollen and honey. 1 food container with at least lKg of milk candy (1 part dry skimmed milk; 6 parts icing sugar; 3 parts honey). (=7 combs plus food container).

90 Obviously any drone brood present on the worker brood combs used must be destroyed. If there are any emerged drones present the combs must be brushed and sieved (an advantage of using brood from reserve colonies is that in May they usually contain no drones). For good drone rearing it is of prime importance to use brood combs of the highest quality, ready to emerge and well covered with bees these should then be given to powerful colonies "boiling over" with bees. The nucleus receives a ripe queen cell from a breeding line. If such a queen cell is not available at the time when the nucleus is made up a "wild" queen cell from a pure breeding line (in an emergency the nucleus can be given the facility to raise its own queen to start with). The remain- der of the drone colonies will be made up in the same way as soon as more drone combs are ready to emerge. Drone nuclei are taken without supers to the Mating Station. The smallest size should be a 10 comb nucleus box. 10-12 combs well insulated on all sides have proved best. At the start of the breeding period, these can be reduced to the desired number of frames by means of a division board and packing. After being kept three days in the cellar, the nucleus can be set up at one side of the apiary, with an excluder or a drone trap over the flight hole as appropriate or it can be taken directly from the cellar to the Mating Station. If all the preparations go smoothly, all the drone stocks can be transported to the Mating Station at the same time. As soon as the bees have started comb building, the best drone-brood is obtained in freshly built comb. For this, a wired frame with only a starter is put into the brood nest. This can be done when another drone comb is still at the stage of being filled with eggs. But drone-rearing usually starts before the bees have begun building. The breeder should therefore have at his disposal a good supply of light brown drone comb (i.e. comb in which drones have already been reared once). In order to obtain drones very early for physical character testing we usually insert combs that are half drone cells and half worker cells. Entire drone combs are only employed at a later stage. It is not necessary to erect an expensive bee house at the Mating Station. It is adequate to have the drone colonies in well insulated hives on a simple migratory trailer or transportable stand.

Breeding Time Breeding always requires strict adherence to a time table. It is therefore advisable to co-ordinate the rearing of queens and drones by using a single time table. The start of queen rearing is designated as Day "O". Preceding days are marked as "-" and the following days as "+" By means of this table (page 44) the actions to be taken at the various stages can be easily noted on a calendar. From the time table it can be seen that the queen requires 20 days from the introduction of breeding material to be ready for mating while the drone takes 40 days (from the laying of the egg).

91 The Queen Mating Station should be in operation by 1st June at the latest. Experience shows that the best mating weather is to be expected in the period from 1st to 10th June and, since at this time only a few drones are flying from surrounding apiaries, a much higher proportion of pure matings can be expected. The following time table is recommended for early breeding:- Start of Drone Rearing 25th April. Start of Queen Rearing 16th May. Setting up of Drone Stocks 25th May. Setting up of Mating Nuclei 1st June.

Mating Station Maintenance For as long a time as the Mating Station is in use, requirements for its proper operation must be completely satisfied. This will be attained by:- 1. Continuous feeding of the drone-stocks. The feeder in each stock will always be supplied with milk-candy (1 part dried skimmed milk; 6 parts icing sugar; 3 parts honey). This also ensures the best results for drone rearing in bad weather. 2. Care to maintain the highest colony strength. The performance as drone rearers is less dependent on the absolute size of the colony, than on having the hive filled to bursting point with bees. If colony strength shows signs of declining (as after temporary queenlessness) it should be immediately assisted by giving a comb of emerging brood. 3 Attention to the maintenance of drone numbers. If, at those stations which continue in operation for longer periods, the nucleus method is being worked, each drone nucleus must receive every three weeks a fresh comb of sealed drone brood from the breeder mother. For this purpose, drone combs must be filled with eggs at regular intervals by the breeder mothers left at the home apiary. These will be nursed in the honey chambers of strong stocks between two combs of brood, till the drones are almost ready to emerge. This procedure conserves the breeder-stocks and makes possible the production of a practically unlimited supply of drones. Stocks which have been tended on these principles will retain their drones well into the middle of September, even if they have a young laying queen. If it is desired to stock the Mating Station with drones of a different line during the course of the same Summer, all the drone stocks and nuclei must first be removed to a distant apiary. Flying drones will return again from 5-6 KM distant. The new drone colonies must be isolated for some time before being conveyed to the Mating Station to discourage the importing of stranger drones.

Which Stations are reliable? When natural mating takes place, the result must in every case be moni- tored by an examination of physical characters. Only by this means can it

92 be discovered what actually took place at the mating station and whether performance was what was expected from the Station. A Mating Station can only be considered serviceable if at the very least 75% are faultless pure matings. This result is by no means easy to achieve at Land Stations where "sneaky crossings" have always to be reckoned with (the queen may have mated for example, with seven Station drones and one intruder). If the percentage of complete pure matings is lower, expenses are not covered, as faulty and mixed matings are only fit for use as commercial queens. A breeding strain can be ruined in a very short time at a bad Mating Station either because of not immediately recognised cross matings, or from such a low output of pure mated queens that an effective selection for performance is no longer feasible. Only on islands, or in districts with an area of at least 8Km radius free from alien drones, can Mating Stations be reliable. Only in such places can pure line breeding be carried on (p.82). To test the reliability of mating at such Stations a Cordovan-test should be carried out at least once. The character "Cordovan" (leather coloured) is the result of a mutation which is inherited as a hidden (recessive) character. A Cordovan (cd) queen only produces brown offspring when she is mated with a cd-drone; when mated with a dark ("wild") drone she produces dark bees. Using this mutation, which is quite specific, Station tests can be carried out very precisely under natural conditions. The Mating Station must first be cleared of other stocks and nuclei. cd-Virgin queens are taken to the Station together with a few cd-drone colonies. From the proportion of pure mated queens, and from the ratio of cd-bees : dark-bees from mixed-mated queens, the number of matings with foreign drones as a proportion of the whole can be calculated very accurately. An examination of physical characters is not suited to this purpose, since it cannot distinguish between matings with Carniolan drones from the Mating Station, and stranger Carniolan drones from colonies in the surrounding area. Experience tells us that no opportunity should be missed for finding out exactly what goes on at the Mating Station. An area of at least 4Km radius free from stranger drones, and with a low bee population around it, is needed for a reliable Queen Mating Station. The success of its operation relies very heavily on the strength of the drone force and the time of breeding (early breeding!) Contrary to previous opinion no general conclusion about the effect of land formation (mountains, plains) on performance of a Mating Station seems possible. Up to now the lowland Mating Stations have clearly been held in low esteem only because stranger stocks were generally speaking established in too great numbers, and with too little separation from the Station. In the mountains, the bee population is generally sparser. Security can be further increased by requeening apiaries in the preferred flight direction Figure 35 shows diagrammatically what detrimental influence the drones of a large apiary situated on the edge of the guard ring can have on a Mating Station if it is not adequately supplied with drones.

Most of those Land Mating Stations hitherto in use, which either have not achieved the 4Km radius, or even fall well below it, and are in the middle of an area thickly populated with bees, are useless. Experience shows that in such a situation merely to increase the numbers of resident drones would not suffice to reduce the invaders to a minority (Figure 35). The only practical, economical and sensible course in such a situation, is to forego pure breeding in the strict sense from the outset, and change the Queen Mating Station into a mating apiary; or create a small pure breeding area with the Queen Mating Station at the centre ("Pure Race Breeding"). The results will be far better in every respect than those which hitherto have produced unsuitable Mating Stations of the traditional kind. If the greater part of the stocks in the neighbourhood can be successfully incorporated into the plan for the Queen Mating Station, and are persis- tently requeened from the preferred breeding line, over a number of years

94 a "useless" Queen Mating Station will be transformed into a very productive "Pure Race Breeding Station". Examples of such a development can be produced in plenty. The daughters of a tested Carniolan queen are not likely to mate exclusively with drones of the same degree of selection; but at least they will mate with drones of the same race and similar derivation. The quality of the stocks built up from them will be satisfactory and the displacement of hybrids and the country race by the constant reinforcement from this island of pure breeding will proceed continuously.

95 Chapter IV The Races and Strains of Honey Bees of Central Europe

1. Distribution Two races of bees have been present in Central Europe from time immemorial ;- 1. The Carinthian (Kärntner) bee, Apis mellifera carnica. (In English they are named Carniolan, the district in Austria from which they were first imported.) 2. The Dark Bee, Apis mellifera mellifera (these are usually called "Nigra" from the colour and from the best known breeding strain). For an exhaustive study of bee races, consult G. Goetze. The Honeybee in Natural and Artificial Selection, Part 1 Verlag Parey Hamburg 1964. F. Ruttner : Biogeography and Taxonomy of the Honey bee, Berlin 1988. Carniolan and Mellifera were originally separated from each other in the Alps by a very distinct boundary. The great distribution areas of the two races march together here; the Dark bee advanced from the North and West, the Carniolan from the Danube basin and from the Balkans into the Alpine region. So, to the South and East of the Alpine ridges we find the original settlement region of the Carniolan, to the north of the watershed and westwards that of the Mellifera. (Figure 36). More in detail: Voralberg, Tyrol, Salzburg without Lungau, Upper Austria, with Styrian Ennstal and Western Lower Austria were originally "dark"; the rest of the region East Tyrol, Carinthia, the greater part of Styria, Burgenland and the East and South of Lower Austria were "grey". The Western boundary of the Carniolan range passed through the Puster valley near the Austro-Italian border. There the Carniolan was bounded on the West by the Dark bee, in the South by the yellow Italian bee. The Venetian Alps are predominantly "grey", in the Western Julian Alps and also in Istria pure grey bees are found, whereas the Venetian plain is already an area belonging to the Yellow bees. South Tyrol is a "three land corner" with black, grey and yellow influence. The original bee here might have been the Dark bee, as in the whole of the Western Alps.

96 In the East the boundary of the Carniolan is not so distinct, as the natural barriers are less clearly defined. The bee of Slovakia is clearly a Carniolan with a more or less strong "N"-admixture. In Bohemia and Moravia there is racial mixture, predominantly Carniolan in the South-East, with stronger "N"- influence in the West and North. In Southern Poland (Galicia and Tatra) a distinct Carniolan character has been observed; in central and North Poland are colonies with pure Mellifera (N or dark) characters. In Russia the quite Carnica-like steppe bee of the Ukraine merges quite gradually into the Dark bee of the Northern Forest Zone. The Carniolan bee therefore extends from the South Eastern Alps to the Black Sea and the South Russian steppe. It is the bee of the Danube basin, and the Northern Balkans — an area with an unqualified Continental climate — cold Winter, short Spring, hot Summer. The distribution area of the Mellifera embraces the whole of Western and Northern Europe and the Northern part of Eastern Europe as far as Siberia. The Dark bee shows thereby great adaptability to the equable Atlantic climate in the West, as well as to the undoubtedly continental climate in the East.

97 The smallest distribution area among the European bees is that of the yellow Italian bee. It is restricted to the Italian peninsula from the Alps to Calabria together with Corsica and Sardinia, but excluding Sicily. The vigorous beekeeping activity of the last hundred years has brought about a very great change in these original conditions. Everywhere in Central Europe the Dark bee has given ground under pressure from the Carniolan. There are no longer any entire districts in the whole of Central Europe with pure Dark bees; a long search must be made before such bees can be found in single isolated apiaries. The "local" bee is now an uncontrolled mish-mash of the native Dark bee and the Carniolan. The proportion of the Dark bee depends on how many Carniolan queens have been brought into the particular district in the course of time, and how greatly they have increased there. In remote areas and in the Western Alpenlands (Salzburg, Tyrol, Corarlberg, Switzerland) the local bee still preserves a predominantly "Dark" character. A further element in the racial medley of the local bees of Central Europe is the genetic influence of the Italian Bee. Formerly this was distributed by imports of the pure race, and extended by their increase in a few places, but now it results mainly from the purchase of "Heather Bee" swarms (the "Heather Bee" now includes a very high proportion of Italian in its make up), and through the widespread adoption of the much vaunt- ed racial hybrids (Caucasit, Buckfast) Italian influence is easily recognised by one or two yellow abdominal rings without the need for a tedious inspection procedure. Although the pure Italian bee is very quiet and gentle, its hybrids become exceptionally excitable. The ousting of the Dark bee by the Carniolan is a process that has gone on quickly and seemingly without a pause. In many districts, for example here and there in the Northern Alps, many very uniform Dark bees could still be found just after the 2nd World War; in the same apiaries only Carniolan hybrids are now to be found. To understand this development, we must compare and contrast the qualities of the Carniolan and Nigra bees.

2. Comparison of Carnica and Mellifera What appears to me as the most vital character of the Carniolan is the spring development, which proceeds by leaps and bounds. They winter in comparatively weak stocks, with the usual type of management on not more than 6-8 "Normal" combs (IV.l.) with correspondingly small stores. This small winter colony strength is in part typical of the Carniolan strains which used to be kept but is also a consequence of the method, common for very many years in the German speaking regions of keeping bees in small back-opening hives. Since the Institutes and Commercial Beekeepers mainly work with the storey hive which can be extended as required, Carniolan strains with very large winter colony strengths have developed such as have been known for a long time in Hungary and Rumania. This is

98 a further example of genetic versatility within a race, which makes adapta- tion to a desired type possible by breeding. In Spring however, there is an extraordinarily rapid, even turbulent, build up which can soon fill the honey chamber. Accordingly the Carniolan is the "ideal bee for the early flow" as Brother Adam, a very critical observer, has called it. This unparalleled vitality and capacity for development is attributable to good resistance to disease (for example out- breaks of Foul Brood are very rare in the whole of the Carniolan region). Another consequence of this quality is the lively swarming impulse, which is without doubt more evident in the Carniolan than in other races. But here too a great deal has been achieved by breeding with proper management, which mainly implies the use of large hives: the proven breeding strains do not display any excessive swarming tendency. The Carniolan bee has provided convincing proof of its industry in many productivity tests throughout the world. In most nectar flows it sur- passes all other races, most strikingly in the early flow and the red clover of which it can make particularly good use, thanks to its long tongue and foraging energy. Indeed the Carniolan bee — most particularly in its specially long-tongued forms is, according to Professor Goetze and Dr. Bottcher, the only bee suited to the Central European climate that can utilise the red clover for us. The exceptional gentleness and quietness on the combs makes the handling of this bee undeniably pleasant. Her scanty use of propolis and very low inclination to rob are also agreeable traits. Finally, and without doubt, the Carnica of all the races we know has the best sense of direction and least tendency to drift. With the Dark bee the difference in colony strength between the full strength of Summer and Winter is less noticeable, the Spring development takes place more slowly and evenly. It is a bee for the equable climate, and the late flow. It is especially esteemed for the heather and the forest flow of high Summer. The brood nest always stays compact and uniform, its self-sufficiency is dazzling. The ideal type of the Nigra (the Swiss strain of the Dark bee) is the "Hungler" as demonstrated to us by the Swiss breeders. The stock with the tight ball of brood surrounded by belts of pollen and honey. At rough sites, with a meagre flow — largely from the forest — the Dark bee or the local bee should be superior to all imported strains. Coupled with the low breeding rate is the disinclination to swarm, an advantage which it undoubtedly has over all but the most thoroughly selected Camiolans. The Dark bee is often excitable and stingy, many colonies are real "runners" while others remain relatively still, and are quite pleasant to handle. Certainly the Dark bee is hardly ever as quiet and gentle as the Carniolan. The argument between the supporters of these two races — the native Dark bee and the imported Carniolan — dominated the field in Germany for many decades. Nowadays the contest in this form has finally exhausted itself, simply because no one takes the trouble to breed the Dark bee any more and the pure strains are no longer obtainable. Anyone who wants

99 pure Dark bees must turn to France, Norway or Poland. In Germany only bees of strongly marked Carniolan type, or variegated hybrid local bees, are to be found. Yet the argument about the "Best Bee" is far from settled. It has simply switched to Carniolans versus Hybrid, (p. 98 para 3 above).

3. The Incompatibility of the Honey-bee Races. Here we must introduce a topic that no beekeeper can avoid. No one can breed bees just as the fancy takes him. Everyone is tied to the race that dominates his neighbourhood, at least as regards breeding objectives. The ground for this statement is very simple; queens and drones cannot be shut up in a box for mating, they unite only in the freedom of the open air, and their flight range extends over many miles. All the drones that fly from an apiary influence not only the one they belong to, but all the other apiaries in the district. The belief that queens only mate with drones from their own colony has long since been shown to be a fairy tale. If anyone brings stocks of a foreign race to his apiaries, unless it is very well isolated, he will hybridise the entire neighbourhood. On the vexatious results of this we must say even more. In doing so there is no intention of defending the primitive condition. Today a bee should only be recommended if it gives the best economic results. No other consideration — tradition, native origin, colour and the like - can be entertained. One thing is certain; in a single district two races cannot be kept together. If a foreign race, after careful testing has been accepted as the better for that area, the entire region must be turned over to this race. This is what happened in Israel. There since the war, the sting-happy unproductive Palestinian bee has been replaced by the imported Italian bee. Thousands of stocks were systematically requeened. For a full century all kinds of experiments were tried in Germany. Into the province of the native Dark bee Carniolans, Italians, Nigra and Cypriot queens were introduced indiscriminately. The result was an unholy racial mess from which at length the Carniolan bee emerged as the undisputed champion. Without any guidance from higher authority practically all progressive beekeepers in Germany today have gone over to the Carniolan. This bee has succeeded wholly because of its superiority. Under pressure of circumstances and with a great deal of trouble the conditions have been created in Germany such as had been present from the beginning in the greater part of Austria. It was only possible to do this because in Austria and also Jugoslavia the Carniolan had been kept pure and tested strains of these could be obtained from Austria.

4. Cross Breeding The racial hybrid that we can obtain most easily is the Carniolan x Nigra (or the reciprocal cross), at least so long as a pure Nigra strain is available. In one

100 of our own trials, pure queens from a breeding line of the one race were mated with pure drones of the other at a secure Mating Station; the hybrid stocks gave a 30% higher yield than those of the pure Carniolan line. In this test the Nigra line was 30% below the Carniolan. The hybrid colonies were exceeding- ly vigorous in brood production, build up and in overall vitality. They also wintered well. However, they had a drawback which was decisive, they were sting-happy and restless. After several years work with these crosses, we had to abandon them in spite of their excellent output. It is not a simple question of the difficulty of the beekeeper's own work; far more serious are the never ending troubles with neighbours and the difficulty of obtaining consent for setting-up out-apiaries and migration sites. The more populous our country becomes the more we are restricted in the siting of apiaries and sites for migratory beekeeping, and it becomes correspondingly more important that we should pay attention to gentle temper in our bees as the deciding factor in breeding. Unfortunately, most racial crosses incline towards increased stinginess, even when outstandingly gentle races are used for crossing. It must once again be emphasised that these good performances by racial hybrids were only attained in the first hybrid generation. By the third generation the average performance was clearly less than those of the parent lines. This was also true of the further crosses, not to mention the very great variability. In the long run, hybrid queens should therefore only be produced in specialised breeding centres which are in a position to maintain the parent strains in a pure state. The crossing of Caucasian x Carniolan proved even better as regards yield than Carniolan x Nigra, and these hybrids also remained fairly gentle. In the course of time however, it became clear that with these bees in hard winters severe losses had to be accepted because of extreme susceptibility to Nosema. Bilasch (Apiacta 1962/2) had the same experience in Russia. The Italian x Carniolan cross gives very good yields, but is very sting-happy. The reciprocal cross on the other hand is gentle but mediocre in performance. (F.Baumgarten A.D.I.Z. 1967/4). The yields from hybrids between (unselected) Italian and Carniolan at Erlangen were distinctly below the average of the pure-race Carniolan control-stocks. The innumerable attempts at crossing which have been made in the course of time, have not succeeded in discovering racial hybrids which satisfy all requirements and which on a long view were the equals of good Carniolan strains in the conditions of Central Europe. Brother Adam's Buckfast Bee provides an exception. There is no doubt that these bees can be reckoned among the most productive and adaptable bees that we know. Particularly noteworthy is the great colony strength with the corresponding potential for high yields and the low swarming tendency. According to Brother Adam this bee was originally derived from a cross between the Italian and the old English Dark bee (but of the influence of the latter there is certainly nothing to be observed neither in behaviour nor in external physical characters in the present day Buckfast bee).

101 Other imported races, such as the Macedonian and the Anatolian, seem to have been repeatedly added to its make up. It is clearly a question of a complicated hybrid which over a great many years has undergone continuous, rigorous selection by Brother Adam with his great experience and his own natural gifts. (Brother Adam A.D.I.Z. 1978 65-71, 101 - 107). The development of this bee, which proved itself in many countries, is without doubt a splendid breeding achievement. Particularly good results have been reported from districts where the Buckfast queens used for requeening mate with pure Carniolan drones , so that repeated hybridising has taken place. In the lively discussions about the Buckfast bee which have taken place from time to time, a certain disenchantment has crept in, no doubt because extravagant claims have led to unfulfilled expectations. Careful tests carried out quite objectively have not demonstrated any superiority of the Buckfast stocks over Carniolan stocks — not even where it was most expected, with a management system of large colonies in a very good honey year. (Dr. V. Maul A.D.I.Z. 1978/5 PP 140-2). The same assessment was made by professional beekeepers with great breeding experience using Buckfast daughter queens either apiary mated or mated with Carniolans. (F. Baumgarten A.D.I.Z. 1978/3 pp 71-77). The cost and trouble of obtaining genuine hybrid breeder queens is also a matter of some consequence for the professional beekeeper. Since therefore it cannot be honestly asserted that the Buckfast bee has an overall superiority of performance, another argument gains greater force. There are no long term plans for hybrid breeding in Germany. Buckfast queens from an authorised dealer, at a suitable price, can only be had in very limited numbers. Therefore "Buckfast" queens are obtained from somewhere or other (usually they are Fl queens mated with Carniolans) and latterly requeening has been carried out with these. Under favourable conditions this has produced useful results for one, or at most two generations. In one of our own investigations apiary mated "Buckfast" queens produced barely 50% of the apiary average, while their sisters, inseminated with selected Carniolan sperm were among the peak producers. The good performance of hybrids is therefore a very transitory phe- nomenon which is soon over and done with. Meanwhile uncontrolled hybridisation, with all its harmful consequences, has occurred. This method is neither genuine hybrid breeding in a technical sense, nor is it a breeding system which will enable the beekeeper to live at peace with his neighbours. Looked at from the interests of the beekeeping industry as a whole, we must conclude that in Germany predominantly harmful results are to be expected from hybrid breeding even with Buckfast bees, which in themselves are very good. (F. Ruttner, A.D.I.Z. 1978 97-101, Der Imkerfreund 1980, pp 247 - 254). On the other hand, the scheme of pure Carniolan breeding offers to all levels of beekeeping many and various possibilities of development. It is based on the active co-operation of many concerned beekeepers free from commercial considerations. Each one has

102 the scope to display his own initiative in the field of breeding and to endeavour, in co-operation with others, to develop a bee which suits his own purpose. It is the aim of this book to encourage this line of work, and to guide it along the right road. The system put forward for breeding of Carniolans has been practiced for decades, and as has been shown, it brings advantages for every level of beekeeping — for the quite passive ("let alone") beekeeper, for the average beekeeper who practices simple queen renewal, as well as for the genuine breeder with his own Performance Selection and Mating Control. This approach will prevent the beekeeping industry in Germany from sinking for a second time into the chaos of uncontrolled cross breeding, such as was formerly the case.

5. How Do Carnica and Mellifera Differ from Each Other? Most beekeepers in Germany are accustomed to judging races solely by the colour of the abdomen. Bees with yellow abdominal rings are "hybrids". This only applies to crossing with yellow races, e.g. Italian or Buckfast bees, which, North of the Alps, only play a distinct role here and there. Carniolan and Mellifera bees both have dark bodies, both occasionally have brown spots or a single ring. The body colour is consequently unreli- able as a distinguishing character. Nonetheless it is easy with some practice, to distinguish the two races by sight. The Carniolan (Figure 37a) is a slender bee. The abdomen is pointed like a torpedo. The arrangement of hairs is the most striking feature; the tomenta are broad and thick, the overhairs are short. This makes the bee look grey and furry, especially when young. "Grey" is the general impression that observation at the flight hole, or of an occupied comb, produces. The Mellifera is larger and more burly, the abdomen is broad. (Figure 37b). The tomenta are narrow and scanty, the overhairs not so abundant, but long. Consequently the dark body colour shines through to a greater extent and the bee appears dark with a characteristic velvety sheen. This is also the general impression from inspection of a stock or observation at the flight hole. (IV. 2.) The recognition of hybrids is more difficult than distinguishing the pure races. The detection of a foreign racial influence from a "sneaky" cross- mating which might have very undesirable effects on future breeding is especially difficult. Very accurate measurements of the wing veins must therefore be carried out, in particular the Cubital Index permits very precise judgements to be made. Details of this in Chapter V, "Assessment of Physical Characters" (p. 111). For practical men, Guido Sklenar has laid down a very memorable description of the Carniolan — Quiet, Gentle, Grey! In fact the shortest def- inition of the most important identification marks for a first assessment. Hybrids are always more nervous and frequently more aggressive than the pure Carniolan. The tomenta are often narrower.

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6. Breeding Strains Experienced breeders in Austria and Slovenia started many decades ago to increase the best of their Carniolan stocks and to replace the bad queens in their apiaries with good ones. By this simple, but long practised selection, many apiaries of uniform, good performance have been created. Some of these breeding strains have on the basis of repeated trials achieved wide distribution.

104 The Sklenar strain (breeder, Guido Sklenar, 1871 - 1953) is derived from the so-called Lower Austrian bee, concerning which Alfonsus wrote a good deal at one time. It is not a separate race in the zoological sense, but a sub-form of the Carniolan from the lowlands, (vine growing climate). The Lower Austrian bee cannot be distinguished with certainty from the Mountain Carniolan by its outward characters — its hair is just as thick and short, the mean Cubital Index of the colonies lies between 2.4 and 3.0, the proboscis is long, the bee is also very gentle and does not propolise. The body colour is generally of a slightly lighter tint and the leather coloured rings and spots are slightly more common than among the bees of Carinthia and Styria. For many years in beekeeping circles, there has been vigorous con- tention about the brown rings of the worker bees in many Carniolan stocks, and Guido Sklenar especially was in a very difficult position throughout his life on this account. Too deeply rooted is the belief that all yellow and brown colour markings derive from crossing with Italian bees, and therefore every colony of this kind ought to be excluded from increase. Sklenar on the other hand had from the beginning rejected "Breeding by colour" and had adopted the perfectly correct viewpoint that brown body marking is not a racial character. Throughout the Carniolan region — even in Jugoslavia — alongside completely dark stocks, there are others, a distinct percentage of whose bees have a brown ring; moreover, the brown ring of the Carniolan is clearly distinguishable from the yellow of Italians. In colour, the Carniolan is by nature just as little uniform as any other race of bees. The leather brown spots of the Carniolan are therefore not signs of hybridisation, and their appearance need not be so strictly evaluated as deviation in the Cubital Index or in hair length. A uniformly dark stock is perhaps more pleasing to our eyes, but it is neither of "purer race" nor, on account of the colour, more productive than another. To be sure as great a uniformity as possible is desirable but this uniformity must show itself above all in the performance of its descendants. There would certainly be no objection to breeding for colour as such, if it were not so difficult to select for colour and performance at the same time. In this matter it is best after all to pursue a reasonable middle course. In a predominantly dark line — assuming that they gave equal performance — dark bees could be preferred as breeding stocks. In a line which has produced' dark bees for generations it is advisable to exclude from reproduction stocks which show much larger brown marks since there must clearly be a suspicion of outside mating. On the other hand it would be a great mistake to exclude a line with good performance simply because it showed brown marks. One thing should always be kept in mind. The development of a line that gives a performance which is well above average, and truly heritable, is a difficult undertaking and one which requires a great deal of time. On the other hand the selection of a pure dark line from any Carniolan strain you

105 choose can be carried out without much trouble within a reasonably short time. In contrast unmistakable yellow rings never appear in the pure Carniolan. They are to be rated signs of hybridising with a yellow race. The Troiseck strain (breeder Jakob Wrisnig 1875 - 1952) comes from the Miirz Valley, Upper Styria and is named after the Mating Station, Troiseck, above Kindberg. This bee is a thoroughbred mountain Carniolan, with all the physical characters and good qualities of its race. Like the Sklenar bee, she has been tested over many decades in the most diverse climatic conditions in many parts of the world. Since 1948, several breeding lines have been developed from colonies of this strain at the Bee Unit at Lunz am See (Lower Austria) through continuous selection at the testing apiaries using the principles set out in this book. The names of these lines ("1012" "1075" "07" or "Wintersbach") are well known in breeding circles. On account of the selection work carried out at Lunz for more than 30 years, this bee is generally referred to nowadays as the "Lunz Strain". The Peschetz strain which has had wider distribution in Germany than any other, comes, according to the breeders, from East Tyrol, and therefore from the most Westerly corner of the Carniolan region in Austria. The Mating Station at Horitzen in Carinthia where the strain was first developed ceased to exist in 1945, but it was possible to keep the strain pure at other stations thanks to the use of the Island Mating Stations, in Schleswig-Holstein for example. The question is always being asked which of these strains is the best? It is quite impossible to give a definite answer to this question. As everyone knows the "Best Bee" does not exist. At the most we can speak of bees that are especially well adapted to particular forage conditions. It so happens, for example, that neither the Troiseck strain nor the Sklenar strain are themselves completely uniform in behaviour. Among the Troisecks we found very prolific colonies, some that had a decided "Hungler" character, swarming and quite non-swarming colonies, early breeders and late breeders. What character the bee possesses depends on the particular line, or alternatively, on the Breeder. In general the different Carniolan strains cannot be distinguished from each other with certainty by external body characters. The names are used simply to indicate the source from which they were derived. As far as yield is concerned, one hears quite frequently from beekeepers in different districts with an all round nectar flow that one strain has performed just as well as another, and that none could be said to be definitely superior. Where there is a markedly early nectar flow in Austria the Sklenar is often the favourite, as they usually raise brood very early and so fill their boxes in good time. This trait can be a decided drawback in areas dependent on the Forest flow, as it can lead to untimely swarm preparations. In these districts the Troiseck strain is nearly always preferred. Our two well tried Carniolan strains therefore complement each other in a very fortunate

106 manner, and there is no point in engaging in useless arguments about them. In case of doubt any beekeeper can try out both strains in his own apiary. The Sklenar, Troiseck and Peschetz strains are not of course the only Carniolan bees worth breeding, but they are the oldest breeding strains. Efforts have recently been made in Carinthia to select new strains from the native bees there. Warning must be given against "Farmers Hives" of "Grey Carniolan Alpine bees" as described by the dealers. These are Carniolans, selected simply because they swarm freely, quite unfit for a modern system of apiculture, only fit to be used as "bee flesh". It was Jan Strgar (1887 - 1957) who rendered the greatest service to the breeding of the Carniolan bee in Stovenia. Strgar practised intensive selection in conjunction with Mating Stations and created a pure-breed district in the valley of the Bistrica in Upper Carniola. The Slovenian Carniolans selected by Jan Strgar and bred nowadays by his successors are exported to many Countries; just like the Austrian Carniolans. Another Slovenian Carniolan strain from the breeder A. Bukovsek of Gollobrdo near Medwode has proved very good in various parts of West Germany and Switzerland. There is no doubt that in other parts of the wide distribution range of the Carniolan bee there are more, naturally occurring, local strains (Ecotypes) which so far have been scarcely influenced by breeding, and whose attributes have not yet been investigated in detail. Rumania provides an exception, where strains of bees from different districts are being compared, and the most suitable are undergoing breeding selection. Between 1966-8, queens were procured from nine different places in South East Europe, (Austria, CSSSR, Hungary, Jugoslavia, Greece, Rumania) and with the support of the German B.K.A. and the German Research Association were carefully tested at 6 testing stations in Austria and West Germany. By this means it was shown that the Alpine Carniolan from Austria and Slovenia is the strain best suited to the conditions of Central Europe. At all Stations the Austrian strain Troiseck-Lunz came out best. The Macedonian bee, which, according to the latest research must be regarded as a separate race, also shows close affinity to the Carnica group. By combination of strains from two different original habitats within the distribution range of the Carniolan, an increase of yield, as high as that from a good racial hybrid, can be obtained. (F. Baumgarten A.D.I.Z. 1967). Thus the good effects of hybrid breeding without its drawbacks can be readily obtained within the framework of a single race. As regards the Dark bee, in Austria only the Braunelle strain (Breeder Fritz Niederwieser, Mayrhofen in Zillertal) may be mentioned as having preserved, up to the present, the genuine characters and behaviour pat- terns of this race. The "Braunelle" stems from a high up, farm-type apiary remote in the Upper Zillertal. The bee is very dark, with a broad abdomen

107 (Figure 37b) and is in selected lines a thoroughly pleasant bee to keep. The compact arrangement of the brood nest, and the habit of maintaining a good reserve of stores deserve special mention. In external characters it conforms to the "standard" typical of the Mellifera race. For some years now the Regional Association for Bee Breeding in the Tyrol has actively supported the breeding of this strain. By setting up secure Mating Stations with a bee free area of up to 12Km (for example the "Hinterautal" mating Station at Karwendel) reliable matings and a broader base for selection were created. The bee officially described as the "Country bee" of Switzerland is uniformly dark in colour, but the wing veins and hairs clearly show it to be a hybrid. This is probably to be explained by the extensive hybridisation between the local Dark bee and the Carniolan stocks which were imported in large numbers at the turn of the century. After a decree banning further imports and ordering Race Breeding, a general selection towards "dark" forms took place. The best known breeding strain in Switzerland is the Nigra (breeder F. Kreyenbiihl) which has given its name among German speakers to the entire race. Introduced into Germany by Professor Zander, it achieved in the course of time wide distribution. Whether its later collapse resulted from unsatisfactory conditions at the Mating Stations (in Switzerland as well as in Germany) or from deficient adaptibility to present day foraging conditions is impossible to say. From the point of view of breeding it would be regrettable if this bee were to vanish for ever, since at present we do not know whether it might still have a part to play in a future breeding programme. But it might be difficult to preserve it for this possibly distant goal. Thanks to Professor Kobel the "Swiss Country Race" is latterly being bred at a few good Mating Stations under better conditions than before.

7. Other races of significance for breeding The Italian bee (Apis mellifera ligustica) was the first foreign race to be imported into Central Europe. As is well known, Dzierzon kept it in his apiary for many years. In spite of many excellent qualities this bee has never become properly naturalised North of the Alps. Externally, the Italian is very similar to the Carniolan, especially the Cubital Index (2.2-2.8), proboscis length (6.5mm to 6.8mm), hair length (short) and Tomenta (broad). The most striking difference is the yellow colouration of the first three abdominal segments, in drones as well as worker bees, and the rather smaller body size (width of the worker cell 5.25mm compared with 5.50mm of the Carniolan). The hair of both workers and drones is yellowish. The Italian might be described as the "Blonde Cousin" of the Carniolan. Biological differences between the two races are more pronounced. Adapted to the mild wet Winter of the Mediterranean climate, it goes on

108 breeding well into the Winter, and so overwinters with great colony strength and begins raising brood again very early in the Spring. In spite of this,(or possibly because of it), Spring build-up is, in our conditions, much slower than with the Carniolan. Inclination to swarming is generally very weak. In gentleness, quietness on the combs and low use of propolis it resembles the Carniolan; but through its great tendency for drifting and its propensity to robbing it is sharply distinguished from the two Dark Races of Europe. If there is only a single Italian stock in an apiary where there is any danger of robbing, you can be sure that the yellow scouts will be first on the spot. Because of their strong tendency to drift, Italian stocks must be kept in single hives, and not in bee houses. Overwintering Italian bees in Central Europe is more troublesome than for the Carniolan and Mellifera races which are adapted to the conditions. Substantial Winter losses, especially in unfavourable years and locations are not uncommon. Nonetheless, by means of suitable management (very strong stocks!) and selection for years on end, successful beekeeping with pure Italians has been achieved in the far North (Norway and Finland) — an impressive example of the speedy adaptability of the honeybee. By contrast, in the Northern States of the U.S.A. and in Canada, the stocks which are nearly all Italian, are destroyed after the harvest and replaced by package bees from California and Florida in the Spring. In recent years successful attempts have been made, to overwinter bee colonies in these regions by using hardier races, especially the Carniolan. The Caucasian Bee (Apis mellifera caucasica) famous for its long tongue (proboscis) which averages 6.8mm to 7.00mm has often been tried in Central Europe. Outwardly it looks very much like the Carniolan on account of its thick grey hairs, though the shade is lead-grey (compared with the brown-grey of the Carniolan). The thoracic hairs of the drone are deep black however and the wing venation differs strongly, the Cubital Index of the workers is on average about 2.2, mid-way between the Carniolan and Mellifera, but the value varies considerably. Most remarkable is the frequently enormous use of propolis, with which in Winter the bees reduce the entrance of the hive to a tiny hole. The Caucasian is one of the gentler races of bees, though not quite so gentle as the Carniolan. This bee also is very prone to drifting and so cannot be kept in a bee house. Opinion about the economic value of this bee varies, but mostly it is not very favourable. The strongest argument against our use of this bee in our region appears to be its extreme susceptibility to nosema. In bad Winters it is always the Caucasian stocks that suffer most. The economic value of another dark race, the Sicilian bee (A. m. sicula), has been explored by Maryan Alber, and also for a time by F. Baumgarten A.D.I.Z. 1978/3 PP 71-77). In physical characters it lies somewhat between the Carniolan and the Tellian (North African) bee. In behaviour it displays "African" traits, as for example by constructing 100-200 queen cells at swarming time.

109 Bees have occasionally been imported from Anatolia. They are very yellow, and markedly aggressive; they appear to be not a separate race but a mixture of near-Asiatic, Caucasian and Balkan elements. The few that have been imported up to now in Central Europe have proved very disappoint- ing.

110 Chapter V The Assessment of the Physical Characters An examination of the physical characters of the bee is carried out at various stages of the queen's life; once on the young queen's progeny, as a first check on her suitability (Pre selection) and again when the queen is at least one year old, as a component part of the main selection procedure. Young pure-bred queens for which proof of performance is not yet available, can nonetheless undergo a preliminary selection. Its purpose is to test the lineage recorded in the Breeding Certificate, by an examination of the physical characters of her early offspring. By this means mismatings at the Mating Station, or breakdowns resulting from earlier crossings, can be brought to light and faulty queens eliminated in good time. The preliminary test is particularly suitable where purchasers require guaranteed pure mated queens. It is also valuable for assessing future breeder-mothers in one's own apiary, thus avoiding the danger of having to exclude a stock after two years testing because of an unfavourable (character) report. The character assessment can be made as soon as the first worker bees emerge using the same methods as for the main test. It only requires 30 bees for the examination. For the pre-conditions which must be observed before undertaking selection see p. 10 (end of the Introduction).

1. Sampling The examination of a single bee will not serve to create a picture of the condition of a whole colony. A larger number must be obtained. A selection test requires from 50 to 100 workers or drones. Care must be taken to ensure that they are all the offspring of the hive-mother which is being tested. For the worker-bee sample, young bees are taken from the brood combs, never bees from the flight hole or from an outer comb, since there is always the possibility that bees from another colony may be present among the flying bees. Drones must, without exception, be allowed to emerge in the honey chamber and taken from there. Killing the bees for the test cannot be avoided. This is most easily done by placing them in a box or cage (without food!) in the frozen food compartment of a refrigerator for 24 hours, or by sulphuring them. (V.l.)

Ill 2. The Instruments The simplest method for the measurement of the Cubital Index (C.I.) is by means of a Thread Counter, with magnification X10 (see p.119). Measurements can be made more accurately and more conveniently with a microscope, or a stand-mounted magnifying glass with a measuring eye piece in which is inserted a special measuring disc (p.122), or the measurement of the veins can be taken from the image of the wings projected on a wall from an ordinary slide projector. The last two methods are comparable in degree of accuracy. Work with the microscope, when one works alone is quicker, more comfortable and independent of the time of day. Since the method used will depend on the apparatus available all three procedures will be described. In addition the following will also be required for the examination:- 1 hand lens or watchmaker's eye glass x 8, or x 10 1 pair watchmaker's pointed forceps (V/2.) 1 pair fine scissors Some small glass dishes Industrial alcohol Sellotape Insect pins Glass slides 5cm x 20cm (for measuring the C.I. with the thread counter or microscope). Glass slide mounts 5cm x 5cm (or 35mm) for use with projector.

3. The Racial Characters Out of the many characters (over 40) which have been employed in the examination of European races of bees a very few have been chosen (four for the worker bees and three for the drones) which are readily identified and which make possible a very reliable differentiation. For examination the bees must be suitably prepared. In particular, the abdomen must be stretched as much as necessary by using forceps. Bees which have been dead for a few days but are not soft give the best results, since when the abdomen is stretched it remains extended. With freshly killed bees the examination can be made easier by the insertion of an insect pin lengthwise through the thorax and abdomen. Old dried up spec- imens should be softened for 24 hours before the examination in a humidity chamber (a thoroughly soaked piece of insulating board under a cover).

The Physical Characters of the Worker Bees a) Colour Markings on the Abdomen. The colouration of the plates of the abdomen has been given close attention by beekeepers from time immemorial. The great majority of the bees of both the Dark and Carniolan races have uniformly dark colouration of all the abdominal rings. (Figure 38 "O" ; O = ohne, "without"). The second

112 ring of many bees shows leather brown spots (Figure 36 E, (E = Ecke,corner) ("Side spots"). The very frequent small spots which are scarcely visible to the naked eye are classed as "O". In the Carniolan, large spots sometimes meet and form a brown ring (Figure 38 1R). A leather-brown ring should not be interpreted as a sign of hybridisation. In a Carniolan colony not more than 5% of bees with such a brown ring should be present. In the Dark bees, rings are not acceptable. The Italian bee has two bright yellow rings (sometimes only one, sometimes three) and in addition the first segment of the abdomen, the Petiole, is more or less yellow. In the very light Italians ("Golden Bees") 1 to 4 segments are yellow, only the tip of the abdomen is black; with this bee as with the Cyprian bee, the Scutellum is also yellow. A light coloured ring in the Carniolan will only rarely be a sign of hybridisation. In addition to consideration of the colour tint (whether yellow or brownish) a look at the drones (see p. 135) is sufficient to decide whether they are hybrid or not. In the assessment of physical characters, not too much importance should be attached to colour markings, at least on "E" ; the original Carniolan is not at all uniform in the colour of the exoskeleton. Everywhere in its home territory, alongside wholely dark colonies (only 20% of Austrian Carniolan stocks!) there are those in which some of the bees have a brown abdominal segment or large spots (Ruttner, 1969). This is certainly not an indication of hybridisation. As mentioned earlier, the colour-tone is duller and darker than in the neighbouring yellow Italian race. Furthermore, the drones of Italian hybrids have yellow rings, while in the Carniolan bee the colour markings of the males are reduced to "islands" (p.136). The colour of the exoskeleton is only of significance as a distinguishing character for the purpose of racial analysis where there is the possibility of a cross with Italians. In Germany the breeding aim of "uniformly dark" is firmly established because the possible occurrence of Italian hybrids has everywhere to be reckoned with, owing to the despatch of "Heather

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Swarms" (with a great deal of Italian yellow) to all parts of the country and also on account of the Buckfast breed. But because yellow rings are so conspicuous they can also be quickly eliminated. In addition to colour, other characters (excitability; stinginess; swarminess) will frequently reveal the hybrid nature of the stock. Spots and one ring of brownish tinge together with otherwise typical Carniolan characters are not to be judged as signs of hybridisation, (see p. 105). For example, in the Sklenar bee, at least in former times a brown ring was to be found "in every 30 bees" — plainly taken as typical and the founder of this strain always defended these characters proudly against all attacks. External characters are a label for a particular ancestry, nothing more.

b) Pilosity — Hair length. The racial differences in the length of the worker bees' overhair show up most clearly on the 5th tergite, the next to the last; (this is the last tergite which carries a tomentum. Figure 39). The difference between the long haired Dark bee and the short haired Carniolan is very great. Intermediate forms result only from crosses. The examination is carried out as follows;- The abdomen is held in profile against a light background, or against the light, and observed with the aid of a lens. As a standard of comparison the width of the first joint of the foot on the back leg is used, (shown black in Figure 39). If the length of the overhairs is less than the width of the foot, it is classed as "short" (k = kurz), if it is equal to it, the length is classed as "medium" (m). The hair of Mellifera worker bees is appreciably longer than the width of the foot joint, and is classed as "long" (l) The typical hair length of the Carniolan is k or between k and m, that of the Mellifera between m and l. Anyone who has once seen these two forms side by side will never have any difficulty in making an assessment. Carniolan bees with only a touch of Mellifera immediately become long haired, so that this character can be an important indicator of hybridisation.

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Evaluation of hair length can be done better and more simply by a procedure devised by Pfarrer Herold (New Beekeeping School, 5th Ed. 1985. Ehrenwirth - Verlag, Munich). The diameter of frame wire is used as the standard of measurement. (V.3.) It usually corresponds to the width of the foot joint. First, make sure that the selected wire has the correct thickness, by comparing it under the lens with several foot joints. If necessary, the end of the wire must be rubbed down to the correct thickness. If a measuring lens is available (or a micrometer) the end of the wire should be rubbed down to a diameter of 0.35mm, which corresponds to the upper limit of the "short" class; two thirds of the bees of a Carniolan stock must be within this class, while the remaining third can occupy the "m" class (0.35mm to 0.40mm). In a pure Carniolan stock there should be no bees in the "l" class, i.e. with hair length of more than 0.40mm. For measuring, the wire is fixed with adhesive tape to a hand lens, or a watchmakers eye glass with magnification of at least x 6 (Figure 40). The end of the wire is bent inwards, at right angles, exactly at the point of sharpest focus. The overhair on the 5th tergite is held just at the end of the wire. If the overhair is shorter than the thickness of the wire it is classed as k "short", if the same it is "m" (medium) and if longer it is "1". Hair length can also be measured by using a slide projector. The bees are pinned in a row to a soft pad (such as a strip of cork) which takes the place of the slide carrier, and a silhouette in exact profile is projected on the wall. For comparison a frame wire (or an insect needle) of the correct thickness will serve. (E. Braun, Deutsche Bienenwirtsch, 1966, 12). Next to the Cubital Index, the hair length is the most important body character for differentiating between Carniolan from Mellifera.

115 The Hairs of the Tomenta The difference, at once apparent to the naked eye between the "grey" Carniolan and the "dark" Mellifera is mainly produced by the tomenta; the broad very thick bands give the Carniolan a "frosty" appearance (Figure 41F). while the dark background shows through the sparse hairs of the narrow tomenta of the Mellifera so that the entire abdomen of this bee appears dark. It is therefore a character like that of the body colour, which can be assessed with the naked eye. (Figure 41). Often much work can be avoided by making a "pre-selection"; stocks of a Mellifera strain whose bees show very distinct tomenta ("zebras") require no further investigation; they are definitely crosses. Among the local "country" bees the naked eye can quickly tell whether the Mellifera or Carniolan influence is upper- most. On the other hand a faint "dark" influence to the tomenta is fairly difficult to recognise in Carniolans; such an effect usually comes to light only in later generations as a narrowing of the bands. Carniolan — Mellifera hybrids, with a preponderating Carniolan component, often even have particularly broad tomenta. Since at the present time it is usually exclusively a matter of the assessment of the purity of Carniolan rather than Mellifera colonies the character "Tomentum width" has lost much of its value in the assessment of physical characters. In connection with the often unavoidable "snap judgement" the difference between the shapes of the abdomen must be mentioned again. The Carniolan — like the Italian and the Caucasian bees — is a decidedly slim bee, with a pointed torpedo-like abdomen. In contrast, the Black bees abdomen has a much broader termination. These differences are at once apparent even on the alighting board (Figure 37a and b). In accordance with the official rules for physical character assessment, the middle one (II) of the three tomenta, which is on the 4th tergite, is evaluated. The width of the tomentum is compared under a lens with the remaining posterior part of the same tergite. (Figure 41 II). This examination must be made in good light. The point where the tomentum on the 4th

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tergite is widest, is selected. It is always slightly away from the middle line. The Dark bee's tomentum is noticeably narrower than the dark band of the tergite (Figure 41, class "f"). In class "ff" (narrow Carniolan and hybrids) the grey and dark bands are equal. With the broad-banded Carniolan the tomentum, which in this case is particularly thick and light coloured, is twice the width of the dark band, or even broader still. (Class F). Bees in this class also have an "extra band" on the 2nd tergite (Figure 41 la). It is obvious that the tomenta cannot be assessed on old bees with worn and matted hair. c) Cubital Index No other physical character is as important for distinguishing between Carnica and Mellifera as the Cubital Index (C.I.; it is also often referred to

117 as the Wing Index). This is not merely because the two races are very clearly distinguished by this character — which is also true of the characteristic hairiness. The C. I. can also be measured very simply and very accurately and the results of this measurement allow the detection of the slightest crossing with the other race to be made without any uncertainty for both the Carniolan or Mellifera races.

The Forewing of the Bee The veins which traverse the wings of insects serve to strengthen them and to supply nourishment during the period of growth. In the wing of the bee they enclose segments which are given distinguishing names. At the fore-edge of the wing lies the elongated Radial Cell (Figure 42R), under this is a row of three Cubital Cells (I, II. III). Below the Cubital Cells lies the large roughly rectangular Discoidal Cell, (D). The third Cubital Cell (C III) which lies with its broadest side towards the tip of the wing, is of greatest importance for distinguishing between the races. The general shape shows the racial difference very clearly: in the Mellifera race it is broad and compact, in the Carniolan race it is long and narrow. The clearest measurable difference between the two races is the ratio of length 'a' to length 'b' (Figure 42). The Cubital Index (C.I.) is a ratio which indicates how many times length 'a' is longer than length 'b'. Or, otherwise expressed, how many times 'b' will divide into 'a'. Thus a C.I. of 2.0 tells us therefore that 'a' is twice as long as 'b'. In the Mellifera 'a' is only a little longer than 'b', the average value of the index lying between 1.6 and 1.9. (In drones of this race the lengths can be equal so that C.I.= 1.0). In the Carnica race, 'a' is generally more than twice as long as 'b' the average C.I. of a colony lying between 2.4 and 3.0. The measurements are made exactly between the points of intersection. (Figures 42 and 43).

The Measurement of the Cubital Index The bees even in a completely uniform colony show fairly large variation in their values; the highest values of Mellifera bees and the lowest values of Carniolans overlap. If only single bees were examined, false assessment would be possible. Even more than for the other characters, a greater number of bees must be examined for the determination of the C.I.; the average value must be determined and the Mean values of pure Mellifera and pure Carniolan colonies must always lie very far apart. The C.I. can often be estimated simply by gauging with the eye, but experience shows that serious mistakes can result from this and on the whole the beginner will not get useful data by this method. For this reason one should become thoroughly familiar with the measuring procedures from the outset; only later, with a sure eye acquired by experience, can one make use of a rough estimate to get an informed quick assessment. But there is no substitute for actual measurement. Slight crossings with local

118 bees can generally only be identified by accurate measurement of the C.I. and plotting the Distribution Curve. The more strongly the Mellifera element in the local race is displaced by the Carniolan, the more accurately must the racial characters in the breeding strain be examined.

How to Measure the Cubital Index About two teaspoonfuls of industrial alcohol is poured into a small glass dish; to this is added a drop or two of sugar syrup or diluted honey. With a watchmaker's forceps one forewing is taken from each bee and placed in the spirit. (With the soft wings of fresh young bees it is often better to use a small pair of scissors.) The wings should always be taken from the same side of all the bees in one sample, either the right or the left. When all the wings have been collected in the dish of spirit they are taken out with the forceps and laid moist in as neat a row as possible on the prepared glass slide. If the spirit dries out because of the high room temperature before the mounting has been completed it is moistened with a little water. (V.4.) The wings are laid in even, closely spaced lines. Care with the arrangement facilitates and speeds measurement. The wings dry on the plates in a few minutes. The addition of sugar to the spirit causes them to adhere well to the plate, and then they only need to be covered with a second glass plate; the edges of the plates are sealed with adhesive tape. This procedure is for measurement with a microscope. If a slide projector is used, the wings are laid between 35mm glass slide covers. The space must be used economically, (15 wings in one cover) or more can be added if the narrow half of the wing is cut away. If no other instrument is available, a hand lens with at least xlO magnification with a measuring scale or graticule (a "thread counter" with divisions of l/10mm) will serve. The small measuring lens B2005 (from Firma Greiner, Bremen, Niedersachsendamm 71, or Firma Wendelin Huber, Dornbirn, Austria) has proved very suitable. For measuring with a lens, the moistened wings are laid in as neat a row as possible on a glass slide about 20cm long and when dry secured with a strip of sellotape. The scale of the thread counter is laid directly on the taped-down wings. Measuring is done exactly as in the method described below. In consequence of the limited magnification of the thread counter, the measurements are of course not very precise but, with a little practice, sufficient accuracy can be attained to obtain the correct Mean and get an approximate picture of the spread. To increase accuracy, one estimates to an accuracy of half a scale division where the end of the section of vein lies between two points on the scale. The measurements are written down in such a way that by simple division the Index value can be immediately

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calculated. (Since the same numbers are being constantly repeated, it is convenient to have a Division table from which to read off the results thus:-)

a. b. Cubital Index 6 : 2.5 2.4

5 : 2 2.5

6 : 2 3.0 5.5 : 2 2.8

and so on.

The Mean (M) is calculated by adding all the Index values and dividing by the number of bees that have been measured. The addition of long columns can be avoided by multiplying each value of the Index by the number of bees with this value, and calculating the grand total from these part totals:-

Index 1.9 2.0 2.2 2.3 2.4 2.5 2.6 2.8 3.0 Total Number 1 2 2 5 10 16 4 6 4 50 of bees

1 X 1.9 = 1.9 2 X 2.0 = 4.0 2 X 2.2 = 4.4 5 X 2.3 = 11.5 10 X 2.4 = 24.0 16 X 2.5 = 40.0 4 X 2.6 = 10.4 6 X 2.8 = 16.8 4 X 3.0 = 12.0

50 125.0 ÷ 50 =2.5 (Mean value of C.I.)

A measure of the Spread, or Scatter, for the colony under consideration is given by recording in brackets after the Mean value, the highest and lowest individual values observed in the sample, e.g. M = 2.50 (1.9 — 3.0). For measuring with the microscope the following are used:- Magnifying lens on a stand, or a simple microscope with about x40 magnification (higher magnification increases the time taken for measuring; lower is not accurate enough).

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Measuring Eye Piece with a Purpose-built Measuring Scale The measuring scale is based on the facts established by Kausche (Z.f. Bienenforschung 4. 1959 pp 194-198) that the angle formed by length 'a' and length 'b' in all races of the honeybee in both workers and drones, is almost exactly 151 degrees (± 1°). By measuring many colonies of different races, we have abundantly confirmed these findings. Accordingly an angled measuring scale can be constructed whose zero point is laid on the junction of the basal vein of the 3rd Cubital Cell and vein "r" which runs towards the rear edge of the wing (it must be exactly in the middle of the junction point of the veins, marked with a dot in Figure 43). The measurement of the longer section 'a' is from zero downwards and of the shorter section 'b' from zero upwards ( the end points of the measurements are likewise indicated by dots in Figure 43). Therefore, by a single application of the scale, (which must be done with the greatest possible precision!) both sections can be measured in one operation, and the process be considerably shortened. The Index Scale for a measuring microscope is at present obtainable from the Institut fur Bienenkunde 6370 Oberursel/Ts., West Germany.

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The measured values of 'a' and 'b' are set down in separate columns, and in a third column the calculated Index values are entered. The printed record sheets used by the Austrian Selection Masters are printed on the back with spaces for 100 worker bees and 100 drones; on the front is squared graph paper for plotting the distribution curve. (Obtainable from Nö Imkerschule, Wiener Neustadt, Waltherstrasse 6, Austria.) For measuring with a slide projector the distance to the screen, or wall, must be arranged so that a magnification of at least x 40 is obtained. Using a projector with a lens of 10cm F/L a projection distance of 4 metres is sufficient. (In a small room or with a lens of longer F/L the enlargement can be doubled by projecting on to a good mirror and thence on to the opposite wall). Measuring on the wall can be carried out with an angled measuring rod which has the shape of the scale shown in Figure 44. The Index Classes For exact calculation of the Index it is essential to give the values as precisely and accurately as possible. In order to calculate the Mean-values and to show the distribution, bees with similar values are arranged in groups, known as Classes. Breeders generally choose a class interval of 0.2, e.g. (2.0, 2.2, 2.4, 2.6 etc.,)

122 The arrangement in classes gives a distorted picture however since the Index value has been obtained by division (a÷b) and quotients, as is well known, produce not an arithmetic but a geometric progression. An example will make this clean- Assuming that in a Mellifera drone, the sections a and b are of equal length then 50 ÷ 50 = 1.0 In another drone a is as much longer, as b is shorter, so C.I. = 55 ÷ 45 = 1.22 In an extreme case of a Carniolan drone, section a is three times as long as section b, so C.I. = 75 ÷ 25 = 3.00. then, as in the previous example, section a, is again longer, and section b shorter, each by the same amount resulting in a Cubital Index:- C.I. = 80 ÷ 20 = 4.0 By a change in the length of the sections of the same amount, a difference in the Mellifera range of 0.22 is produced, but in the extreme Carniolan case it is 1.00. This is almost a fivefold difference. If the class interval stated above of 0.2 is used, a change of the same amount in the sections produces in the first example a change of one class (from 1.00 to 1.22), but in the second example a change of 5 classes (from 3.0 to 4.0). This is easily understood. In one example the divisor is reduced by 10% (from 50 to 45),in the other example by 20% (from 25 to 20). This shift causes the bees in the lower range of the scale to be compressed into a few classes, while those in the upper range are spread out into many. By organ- ising the classes in this way the distribution curve is artificially "skewed". Since a great deal can be said about the composition of a colony from the shape of the distribution curve, it is very important that the arrangement of classes should not be open to criticism. This is all the more important since many breeders nowadays prefer Carniolan strains with high Cubital Index (M = 3.0 or above). Dr. Dreher has devised a mathematically unobjectionable scale (Z.f. Bienenforschung, 1. 1950. p. 17). He has shown that the division into classes must always be made from the same change in the measured length and not from the change in the Index value. This means that — as shown in the example quoted above — the actual class interval in the lower range is smaller than in the higher range. Dreher takes as the interval between the classes a change of 1/50 in the total length of the sections a + b (the Basal Vein). We ourselves use a class distribution on the same principle, but with a unit of change of 1/60 of the basal vein, consequently:-

Class 6 50.00:50,00 = 1.00 Class 7 51.66:48.34 = 1.07 Class 8 53.33:46.47 = 1.14 Class 9 55.00:45.00 = 1.22 etc....

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With this class distribution (using 1/60 instead of 1/50) better use is made of the precise measurements that can be obtained by using a magnification X40 and more convenient, reliable boundaries between the classes result. The classes are numbered consecutively; 5 classes have to be provided for Cubital Indices less than 1.0 to accommodate the extreme Mellifera drones. Our Index Scale looks as follows:- (Table 2. p. 124)

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Table 3 Example of table used for calculating the Class values of Cubital Index from the measured values of the two sections 'a' and 'b'.

Class 15, for example includes all bees with a C.I. value between 1.86 and 1.99, Class 16, those between 2.0 and 2.15, etc. It is not at all necessary first to calculate the actual value of the C.I. and then to look to see into which class it falls; the Class Value can be determined directly by means of an "Index Table" (Table 3) from which the Class Values can be read quickly and without calculation. The measured values of the long section 'a' are given in the top row; the values of the short section 'b' are in the vertical left hand column. A greater degree of accuracy is necessary in measuring section 'b'; the lengths are therefore read to an accuracy of 1/2 a scale division, which it is possible to do by estimation, while for section 'a' measuring to the nearest whole scale division is accurate enough. If an index of 21: 7.5 is measured, 21 is found in the top row, 7.5 in the left hand column, and the intersection of the two rows gives a Class Value of 20. On the report card, the Class Value for each bee is entered in the third column, opposite the measured values of 'a' and 'b'. The Index Table given here is only an example. Everyone must com- pile a suitable table for his own use, which will differ according to the instruments used, the degree of magnification and the actual values obtained with them. The vein lengths to be measured for drones are appreciably longer than those of the workers, so a separate table must be

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prepared for them. Obviously too, the Dark bee will need a different table from other races. Measurement is quicker still — using the projection method — with Pfarrer Herold's "Index Fan" by means of which the value of the Index can be read off directly. The measure is placed on the image of the wing, so that the vertical line passes through the middle of the point of intersection of 'a' and 'b', and the right hand, sloping boundary line, passes through the end point of section 'b'. The adjustment of the scale to these two end points of section 'b' must be such that the lines pass precisely through the points of intersection of the veins as indicated in Figure 43; otherwise errors of up to two classes can occur. The Class Value is read off on the longer horizontal line to the left. Each class begins with the right hand boundary line of a small diamond and ends just before the left hand boundary; so if the point of intersection of the veins (the left hand limit of section 'a') lies exactly on one of the sloping lines, it has the value of the Class on the left. The scale depicted on Figure 45 is only to be regarded as an illustration; for actual measurements it must be enlarged by about 2.1/2 times (each of the five large divisions of the horizontal baseline should be about 50mm long). The full size Index Fan is obtainable from Landesverband Bayerischen Imker, 85 Nurnberg,

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Schweppermannstrasse 63 (however, only the actual values are given on it).(V 6) Working in pairs is the quicker way, one man to do the measuring, the other to work the apparatus and record the measurements. Using a mirror or a projector with remote control, one can cope quite well by oneself.

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A different instrument for the direct measurement of the Index has been constructed by H. Maag (St. Gallen, Switzerland). Nowadays, measurement with a microscope has been almost universally adopted in selection practice. This procedure is the most accurate and convenient. Even in this there is a method of working for two that is efficient and comfortable for the eyes; one person does the measuring, the other finds the measured values as they are called out, or the Class value on the Index scale and puts a cross in the record chart. (Table 4). It is also advisable when measuring with a projector to enter the measurements on a card of prepared squared paper. (Table 4). Every cross represents a bee. When the measurements are completed, a Distribution Curve has already been constructed in the form of a Histogram. Data can be entered in the same way in the printed form at the back of the official selection form of the D.I.B. All that then remains to be done is the calculation of the Mean (p.120, 132). As soon as all the measurements of a sample have been completed and the Class values obtained from the Index table, the Class values which occur in the sample are arranged in order; in the bottom row the number of bees which belong to each Class is written. (On the reverse of the Austrian printed form, alongside the columns for the individual values, a half column with the heading 'VR' is provided for these groupings). The series of numbers gives the distribution of the Index values of the stock under consideration, which in an average Carniolan stock looks somewhat as follows:-

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The distribution is more impressive if set out in the form of a curve. (Figure 46). The number of bees is plotted in the vertical scale, the Index Classes in the horizontal. The usual representation as a "smoothed" curve gives basically the same kind of representation as a Histogram. To construct it the points are plotted at the middle of the Class divisions (therefore not over 16 or 17, but over the mid point 16.5 etc.) at the level of the respective number of bees. These points are then joined by straight lines. Thus the curve does not hang "in the air"; the two ends are joined to the middle of the next Class, higher or lower as the case may be. Figure 46. For comparison, the same distribution is plotted not with our Class values, but with the usual separation of Classes by equal intervals of 0.2 (Figure 47). From the slope of the curve and its elongated shape at the higher Index values the distortion by this method of representation is clearly perceptible. In the upper region, the Class intervals are too small; in the lower part of the Scale, the Mellifera region, the Class intervals are too coarse. As a result, the curve is shortened on the left and extended on the right. However when the mean is calculated by each method the difference is negligible (2.71 as against 2.70).

Assessment of the C.I. Distribution Curve A good deal of information can be extracted from a Distribution Curve of the Cubital Index. First of all it is important to ascertain whether it covers the range of values for the race in question. The boundary line in this instance is C.I. 2.0, that is to say the left hand boundary of Class 16; this should be indicated on the graph on every occasion. The curve of a Carniolan stock must lie almost entirely to the right of this line; at the very most only 2% of the bees may have Index values below Class 16; in

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general, none should fall below Class 15 (1.86). Conversely, the Index curve of a Mellifera stock should lie to the left of Class 16, and should scarcely ever exceed it. A second peak which indicates a concentration of results in Classes 16 and 17 is always a sign of a slight crossing with another race. This is frequently the only indication of hybridity because the Mean Value of the Cubital Index is still well within the permitted range for Carniolan bees (Figure 48). Of course this second peak also affects the Mean, but not to the extent that the permitted limit would be reached especially if one of the parents has shown a very high value. As Figure 50, (Ruttner 1969) shows, mismating which may be only par- tial, of a queen from a line with very high C.I. can produce a stock whose Mean C.I. lies in the upper part of the C.I. range of this race. In spite of this the cross can be detected by the Second Peak in Class 17. The same effect can be seen in Figure 48, but in this case the whole curve is shifted towards the lower values. By the evaluation of the measurement data from very many Austrian Carniolan colonies which have not been influenced by breeding, and of an equal number of colonies from a breeding strain, it has been shown that in an uncrossed Carniolan stock not more than 15% of the bees fell below Class 18. (Ruttner : Biometric Characterisation of the Austrian Carniolan Bee. Zeitschrift f. Bienenforschung 9/1969, 469-4503). The uniformity of a sample, its spread, can be read from the width of the curve. The narrower and higher the curve, the greater the uniformity. Figure 49 gives the curve of a N-Line (N = Nigra, Black, Mellifera) inbred by artificial insemination, which has a very small spread. About two thirds of all the bees are concentrated in only two classes. In average Carniolan stocks, the distribution curve extends over 7 to 10 classes in the worker bees and over 8 to 12 classes among the drones. Among the latter it is not uncommon to find single Index values lying far from the Mean, away from the main curve. Drones from quite definitely unmixed Carniolan stocks from Carinthia were distributed with some regularity to below Class 11,

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and not rarely single values were to be found even in class 9 or 8. These insects, provided they only crop up sporadically are to be regarded as freaks, and should not therefore be considered in the assessment. The "ideal" Cubital Index for Carniolans (Figure 51a) was obtained from 100 Austrian Carniolan colonies which were uninfluenced by breeding and comprised more than 9000 individual measurements. Single values of the worker bees spread from Class 15 to Class 24. The Mean was 19.80 (equivalent to an Index value of 2.71). The Mean values of individual colonies varied between 2.38 and 3.37, but in 68% of the colonies the mean was between 2.54 and 2.89 so a "typical" Mean value for Carniolan worker bees of 2.60 to 2.90 can be accepted. The curve for the Drones (Figure 51b) constructed in the same way is noticeably broader (Classes 8 to 22). The Mean occurs at 15.89 (Actual Index Value 1.98). "Typical" Carniolan drones (68% of the sample) have a

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Mean Index value between 1.83 and 2.15. As regards the distribution of the individual drones in the lower region of the curve, it should be noted that only 4% of all the drones in the sample fall below Class 11 (below 1.40). In a "typical" Camiolan stock the Index values of the drones should not extend below 1.40 and only a few should fall below 1.50. The Index curve of hybrids is broad, often showing two or three peaks (Figure 48). But if two peaks lie close together in the middle of typical Carniolan territory, it usually has little significance; probably, by measuring a greater number of wings, the two peaks would merge. The Mean, at least for a rough estimate, can be read from the curve. Think of the curve as though cut out of paper and laid over a knitting needle, so that the right and left sides balance each other. The Mean lies on this line. So the presentation of the Index values in the form of a graph allows all the essentials to be inferred from it. In making an assessment one should never proceed in a purely mechanical fashion; one should always have in mind the broad picture of the entire family. A colony which is being considered for selection will also be compared with its parent and sister colonies. Every strongly marked deviation is suspect. The Mean values given in the tables "Racial Characters of the Honeybee" (p. 138) apply to the race under consideration as a whole. Individual breeding strains and breeding lines vary within much narrower limits. In the strains from Carinthia and Styria the Mean values of the Index of the worker bees lie almost invariably between 2.55 and 2.90. Breeding from colonies with a Mean Index value of 2.40 would not be undertaken because they are not "racially typical". As has been said already, the Distribution curve can often provide more information than the Mean value taken alone. The graphs set out in Figure 48 and 50 display relatively high Mean values but in spite of that an outside influence is clearly evident. The calculation of the Mean value (M) is carried out in the same way as that with which we have become acquainted in the simplified measurement using the Thread Counter. The Class value is multiplied by the number of bees per class; the products are added together, and divided by the total number of the bees. The class numbers required for the calculations have already been prepared by grouping the C.I. Distribution (p.122). It is now worth giving some thought to what we have to include in the calculation as the Class value. For example: Class 20 includes all the bees with Index values between 2.75 and 2.99. At 2.74 Class 19 ends, and at

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3.00 Class 21 begins. This is a fairly wide range, and one can envisage that the bees included therein are distributed with reasonable uniformity over it. If the scale is expanded the distribution within a Class will appear somewhat as follows (Figure 52: the points represent the Index Values of individual bees). If the value, which best represents the average of the 20 bees included in Class 20, is being looked for, then neither 20.0 nor 20.9 will be acceptable, but 20.5, the mid-point of the class about which the individual values are grouped. Therefore, in order to calculate the Mean Cubital Index value of the sample, the values of the middle point of the Class must be multiplied by the number of bees in the Class:- 4 x 16.5 = 66.0 b 11 x 17.5 = 192.5 19 x 18.5 = 351.5 23 x 19.5 = 448.5 20 x 20.5 = 410.0 11 x 21.5 = 236.5 10 x 22.5 = 225.0 2 x 23.5 = 47.0

100 1977.0 ÷ 100 = 19.77

It will be observed that in these calculations 100 x 0.5 is included in the total and at the end the total is divided by 100. The calculation can be simplified by taking 0.5 from each class and adding this amount to the final value:- 4 X 16 = = 64 11 X 17 = = 187 19 X 18 = = 342 23 X 19 = = 437 20 X 20 = = 400 11 X 21 = = 231 10 X 22 = = 220 2 X 23 = = 46

100 1927-f-100 = 19.27 + 0.50 19.77

19.77 is the Index Mean value of the sample expressed as a Class value. The Mean value can now be accurately indicated by a vertical line in the Distribution Curve. But it is more usual to think in terms of the Actual Index value rather than the Class value. So in the end, the determined Class value is convert- ed to the Actual Index value. In our scale (p.124) a Class value of 19.0 corresponds to a true C.I. value 2.53. But 19.77 lies much nearer to 20.0 than to 19.0, to be precise 77/100 of

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the Class width above the Value 19. Class 19 comprises the Index range: 2.53 to 2.75. The Class width therefore amounts to 0.22. A Class value of 77/100 of 0.22 = 0.77 x 0.22 = 0.1694 (rounded up to 0.17) 19.00 = 2.53 0.77 = 0.17

19.77 2.70 Actual Mean Value.

Presentation in the form of a Distribution Curve and the calculation of the Mean provide the most important bases for the accurate assessment of the Cubital Index. In the course of time, measuring and calculating procedures have come into use which make possible very speedy work indeed. Once the necessary equipment has been procured a precise determination of the Index can be made with scarcely more demand on time than that for the rough estimate with a lens. Yet the procedure is so simple that amateur breeders trained as Selection Masters can carry it out faultlessly after only brief instruction. In addition to the Cubital Index, other wing vein characters have been proposed for distinguishing races — Radial Index, Precubital Index, Hantel Index, Discoidal shift; yet none of these characters tells any more or even as much about race membership as the Cubital Index, whose value for identifying races has been demonstrated in thousands of samples. More about the other wing-vein characters can be found in the work of G. Goetze; "The significance of the wing veins in the assessment of honey bees for breeding purposes". (Zeitschrift für Bienenforschung Band 4 (1959) pp 141 - 148) (V 5).

Physical Characters of Drones Drones have no tomenta. Their overhairs are very long and difficult to assess. Instead the colour of the hair is noted. Body markings and Cubital Index are investigated in the same way as for the worker bees.

a) Body Markings As with the workers, the colour markings appear mainly on the 2nd abdominal segment. To casual observation this particularly broad tergite appears to be the first. The true first tergite, the Petiole (Figure 53), of the drone is very small and hidden by hair. The visible segments all show a light "saddle stripe" on the rear edge. In the Carniolan this is a brassy- bronze yellow; in the Mellifera it is more or less dark (in the very dark Nigra form it is quite black and no longer visible). The boundaries of these coloured edgings is very uncertain and variable, so that in modern practice they are no longer assessed. The colour markings of the drone which are used for evaluation lie on the second abdominal segment nearer the thorax. In their weakest form,

135

they appear as tiny brown spots, the so called "little islands" recorded as "i". The "little islands" appear in various shapes (Figure 53): a. As small round spots towards the sides near the spiracles (ii), not uncommon in the Carniolan drone, but not easy to find because of the thick hair: b. As a broadening of the saddle stripes (is). According to Dr. Dreher, this form occurs especially in the Mellifera drone. c. As a combination of "ii" and "is" (iis) Dr. Dreher, (Bienenwirtschaft 1961 H 12) says this can be presumed to indicate slight Italian influence. d. "Large Islands" separated from each other only by a dark narrow bridge, denoted by "I". If these fuse together in the middle it produces a Yellow Ring (1R). Usually it occurs as a somewhat lighter dark stripe as compared with the saddle stripe. By further lightening the following tergites may also show yellow rings (2R - 3R). It was formerly thought that the brown rings on worker bees were inherited from drones with small "islands" (ii); on this account during research into physical characters special attention was paid to them V. Maul (Kirchhain) has shown however, that this correlation does not exist. As with worker bees, so with drones, only a yellow colouring of greater extent (I and R) is to be judged as a sign of hybridisation.

b. Hair colour Dr. Goetze's colour scale (V 6) reproduces the colour grades to be found in the European races of bees. The Carniolan drones are 'grey', more often 'mud grey' than 'sand grey'. Not infrequently, especially in the older drones the colour lies between mud grey and rust brown, but rust brown drones are certainly not found among pure Carniolans. "Brown" is the hair colour of the Mellifera drone. Very often rust brown and coffee brown are found together in the same stock. "Black" is found — rarely however — in the extremely dark Nigra form. Caucasian drones are also black haired, so far as I have been able to tell up to now.

136

To decide on the hair colour; the drone is placed between the two areas of one of the colour types, with the underside to the left (i.e. towards the lighter shade) and the back towards the darker shade on the right. The drone's hairs correspond to a colour shade when they do not stand out from the background. The symbols for the colours are; gr. = grey, br = brown, sch. = black, g.(gelb) = yellow. c. Cubital Index The Cubital Index of the drone is measured in exactly the same way as that of the worker. Since the wings are bigger, both vein segments are longer. But since section b is increased to a greater extent than section a, the drones Index is regularly lower by several tenths than that of worker bees from the same strain. This must be kept in mind when the computed C.I. values are being assessed. The Mean value for Carniolan drones is about 2.0 and should not be lower than 1.8. That of the Mellifera drones is between 1.2 and 1.5 (p.139).

4. Interpretation of the Data Whether the physical characters taken together correspond to the race under consideration, or whether there are any signs of hybridisation must be determined from an examination of a sample of bees from the intended breeding stock. For this purpose the determined values are so arranged that an easy comparison can be made on a basis of 100 bees per sample (if 50 bees have been examined, the numbers of bees in each class is multiplied by 2). It is the practice in Germany to record the characters always in the same order as on the selection form. The selection form of a breeding colony looks somewhat as follows;-

Workers Body Length of Tomenta Cubital Index markings overhairs Class O/e E R k m l F ff f Carnica (%) 88 12 - 100 - - 90 10 - Ø=2.7(1.9-3.4) Mellifera (%) 100 ------10 90 - 12 88 Ø =1,7(1.2 - 2.1)

Drones Body Hair colour Cubital Index

markings Class O/i I R gr. ge. br. schw Carnica (%) 92 8 - 100 - - - Ø =2.1(1.6-3.0) Mellifera (%) 100 - - - - - 100 - Ø =1/3(1.0-1.6)

The headings for the Classes are often omitted, but as the sequence is always the same, to which characters the percentages refer can always be found by the use of this form. The "Race Standard" sets the highest allowable percentage for the separate character classes which may be reached by racially typical workers

137

and drones. In the Breeding Regulations of the German B.K.A. (D.l.B.) (1986) the following standards were prescribed for the Camica and Mellifera races:-

Workers Body Length of Tomenta Cubital Index Markings overhairs Class O/e E R k m l F ff f Carnica (%) 100 30 - 100 30 - 100 50 - Ø over 2.5 single bees over 1.8 Mellifera (%) 100 80 - - 70 100 - 70 100 Ø under 1.9 single bees under 2.2 Drones Body markings Hair colour Cubital Index Class O/I I R gr ge br schw Carnica (%) 100 10 - 100 20 - - Ø Over 1.8 Mellifera (%) 100 - - - - 100 50 Ø under 1.5

Deviations of less than 4 per cent (= 2 bees out of 50) are disregarded (this also applies to the occurrence of single rings in the Carniolans).

5. The Physical Characters of Races of the Honey Bee SUMMARY IN TABULAR FORM

Geographical Mellifera = N Carnica = K Ligustica = L Race Dark Bee (Nigra) Carniolan Bee Italian Bee

Range West, North and South Eastern Italian North central - Alps; Peninsula Europe North Balkans Northern Russia & Danube Basin

Appearance Large, broad, Medium size, Medium size,slim short limbs slim and long and long limbs limbs

Abdominal Dark (sometimes Dark sometimes 1-3 Distinct Body Colour small brown leather brown yellow rings Workers spots) spots and 1 ring Extent of Yellow (not a sign of colour variable hybridisation)

Drones Commonly small "0i" Never R

Tomenta Narrow, sparse Broad, con- Broad, yellowish (4th Abdominal FI*)=0.5-1.5 spicuous, grey. FI*)=1.5-3.0

138

Geographical Race Mellifera Carnica Ligustica

Tongue reach =Rt R1 + 0.2mm Rl + 0.2 Rl + 0.2

Overhair, Long. Short Short 5th abdominal (0.4 - 0.6mm) (0.25 - 0.35mm) tergite dense

Hair colour Brown - Black Pure grey to Yellowish Drones Browny-grey

Cubital Index Ø 1.5 - 1. 9 2.4-3.0 2.0-2.7 Workers (typically 1.7) (typically - 2.7)

Drones 1.0 - 1.5 1.8-2.3 1.6 - 2.0 (typically 2.0)

Proboscis length Ø Short = Long = Long = (anatomical) =R1 5.8-6.2mm 6.4-6.8 mm 6.4 - 6.7mm Mean Values

Wax mirrors on Rear edge Rear edge Rear edge the 5th sternite straight curved curved

other Late breeder Early breeder Early breeder attributes Not swarmy good brooder + Not swarmy (The Heide biene More inclined is an exception to swarming +

"Stingy" Sweet tempered Sweet Tempered

Less accurate "Homing" Not homing well homing accurately

Winter hardy Winter hardy Over wintering in large in relatively often difficult colonies small colonies with very meagre stores

excitable Very calm Calm

not so quiet on Quiet on comb Quiet on combs combs

Not effective on "Effective on "Effective on red clover. red clover" red clover"

*= Tomentum Index (Tomentum width: width of the dark, Tomentum-free strip of the tergite (Figure 41)

139 If the physical characters of workers or drones in a stock do not lie within the limits prescribed for the Standard, they are rated as "inadmissible as regards physical characters" and are debarred thereby from selection. A stock is rated as "fit to breed from" if its own performance and the average performance of related colonies lie above the average of the apiary, its attributes express the Breeding Aim, and the physical characters are typical for the race. Stocks which display very characteristic and uniform physical characters receive the classification "specially uniform and racially typical" Once again, we will summarise the characters of a Carniolan stock that is rated fit to breed from:-.

General Description:- Grey, Gentle, Quiet.

Worker Bees Colour Markings. Breeding aim: pure dark, in accordance with the Breeding Rules of the German B.K.A.; only E but no R permitted. In the "Natural Racial Standard" of the Carniolan bee however brown rings appear in all strains with varying frequency. The evaluation of colour markings is therefore a question of breeding policy and not of racial purity. A sudden appearance of rings in a previously uniformly good line is a pointer to cross matings. Hair Length Preferably 100% 'k' (under 0.35mm) but 30% 'm' is permitted. Tomenta 50 to 100% F. Cubital Index Mean; 2.4 to more than 3.0; in most breeding lines between 2.6 and 2.9. Of the individual bees, 2% at the most may be under 2.0; and no bee should have a value below 1.86 (Class 15). There should be very few bees in Classes 16 and 17 (very gradual slope of the Distribution Curve; at the most, only 15% of the bees below Class 18). A small secondary peak, or even a shoulder in the region of Classes 16,17 is always an indication of a crossing with the genes of local bees. For comparison. The Mean-value of the N-bee lies between 1.5 and 1.9 (mainly between 1.6 and 1.8), the value for individual bees between 1.2 - 2.2 In contrast to the Carniolan, only very few bees should lie above 2.0. The Index value 2.0 (between Classes 15 and 16) is therefore a very important boundary line between the two races, and this should be specially indicated on the Distribution Curves. Class 16 is only sparingly occupied by the pure races. With crosses, a peak occurs here.

Proboscis Length 6.5 - 6.8 mm.

140

Drones Colour Markings If the breeding aim is "Dark"; as few 'i' as possible "I" and "R" wholly inadmissible. Hair Colour Grey to an intermediate shade between mud-grey and rust brown. Cubital Index Mean value, 1.8 to more than 2.0, never below 1.7. The values of individual drones have a wide scatter but the closed distribution curve should never extend below Class 11 (1.40). Single values which lie well away from the Distribution Curve need not however be considered, not even in the calculation of the Mean value. The permissible range of variation of a Carniolan and a Mellifera breeding stock together with the limiting value between the two is illustrated graph- ically once more in Figure 54.

6. Measurement of the Proboscis. In the more precise selection of the bee, measurement of the proboscis also plays a part. Certainly the differences between stocks appear to be very insignificant; they amount to only a few tenths of a millimetre. Short tongued stocks of the Dark race have an average proboscis length of 5.8 - 6.2 mm; long-tongued Carniolan stocks up to 6.(3 or 6.8 mm In practice however, this small measurable difference can lead to very considerable differences in harvest, especially in districts with plenty of Red Clover. As a rule, honey harvests from red clover can only be obtained with long tongued races, such as the Carniolan. In connection with the working of red clover, the proboscis length is of interest, not only to the beekeeper, but also to the seed industry. For this

141

142

reason bee breeders, who are concerned with the Carniolan bee with regard to this very promising raw material, should not carelessly ignore tongue measurement. Measurement of the proboscis can be carried out by two different methods. a) Measurement of the Tongue Reach As the name implies this is not measuring the tongue itself, but of the depth from which a bee can suck from in a tube, or something similar. But since the bee pushes part of its head in as well, the reach is a little greater than the anatomical proboscis length. Different types of "Glossometer" have been recommended, but up to now none has gained general acceptance. Since at present there is not one serviceable apparatus available the beekeeper must rely on making his own. It is important that the diameter of the openings, through which the bee can reach the food provided is exactly 2.0mm. A Capillary-glossometer, which permits simple and accurate reading is recommended by Beemaster John Falkenberg, 2418 Ratzeburg, Schleswig- Holstein (Figure 55 p.142). A row of 2 mm holes bored in a Plexiglas block are filled with dilute honey. Reading must take place after a certain fixed time interval (5 or 10 minutes) to avoid errors resulting from evapo-

143 ration. When the glossometer is installed in a colony, several bees can drink at the same time; values are therefore obtained which lie between the average and the maximum depth reached. As figure 56 shows, the bee places its tongue right against the wall of the tube, in order to reach the edge of the meniscus. In order to avoid errors caused by evaporation Pfarrer Herrold (Neue Inkerschule, 2nd Edition 1972) uses a plastic apparatus filled with sugar candy. The bees can take the food through holes bored in the lid. A sliding scale on the side enables the depth of the tongue reach to be made.

b) Measurement of the Actual Proboscis Length This method allows the proboscis length to be accurately determined for individual bees, and its use is preferred on this account . Dr. Böttcher has briefly described a method (Imkerfreund, 1962, No. 9) for which no apparatus other than a projector is required, and so can be recommended for general use. The bees to be examined are killed with ether or hot water so that their probosces are well stretched out. The bee is taken in the left hand between thumb and forefinger the head downwards and the underside to the right. The proboscis can then be readily examined. The individual parts are shown in Figure 57. During dissection the proboscis must not simply be pulled out by the tongue which would just simply break off. The proboscis is therefore first loosened by the insertion of fine watchmaker's forceps into the soft parts between the lora, and then drawn forward. Then one of the two lora is gripped by the forceps and the whole proboscis is removed with it at the same time. Against a light background, all the external adhering parts can be removed with the help of another pair of forceps, namely lora, maxilla, and remains of the muscles. There then remains the rod of the drinking apparatus — submentum, mentum and glossa. For measuring, the proboscis is mounted between two glass plates. For this a small quantity of gelatine is warmed in a water bath till it is liquid. The proboscis is laid with its posterior, inward-bent side upwards, on a 5 x 5 cm glass slide (or on a glass slide-mount 24x36mm). A drop of liquid gelatine is placed on the cover glass; the cover glass is then carefully turned over, so that the drop is hanging on the underside; and then low- ered slowly on to the proboscis. Should it happen that the proboscis is no longer lying quite straight, it can as a rule, still be put right if it is carefully poked on the cover glass from one side or the other with a needle or a splinter of wood. Each proboscis must be placed separately under a cover glass. In a few minutes the gelatine will set, and measuring can begin. Using an ordinary projector and careful focussing, the proboscis is projected on a light coloured wall, and the whole length from the point of the submentum to the tip of the glossa (flagellum) is measured with a rule. The measurement is even more accurate if the length is laid off with a pair of dividers on 1mm squared paper.

144 The magnification resulting from the projection is ascertained by projecting a reference standard. Transparent 1mm squared paper, which is readily obtainable, is very suitable for this. It is best if the projector is posi- tioned at the start so that a particular enlargement, e.g. x 40 is obtained, lcm of the 1mm squared paper in the projector must appear as 40cm on the wall. This is easily obtained by moving the apparatus. If the projection of the proboscis on the wall produces a measurement for example of 262mm then the actual length is 262 ÷ 40 = 6.54mm. Ultimately it is possible, to produce a scale by which one can read off the actual proboscis length directly without calculation. Proboscis length is of course only one factor that influences foraging performance from intricately constructed flowers (e.g. the Leguminosae) Certainly behavioural characters such as aptitude, learning ability etc., play a part. Thus it has been reported that the Caucasian bee did not bring in the expected harvest from the red clover, although it possesses the longest proboscis of any race. Mackensen and Nye (Journal of Apicultural Research, 1965 - 70) were able to show, that by selection according to flying frequency (determined from the frequency of the pollen loads), a bee could be obtained fairly quickly which was specialised to a high degree for a particular nectar flow (e.g.the "Lucerne Bee"). That a "red clover bee" must not only be adept at working this flower, but must have a long tongue as well, goes without saying. The physical characters here cited — Body Colour, Body Hairs, Cubital Index and proboscis length — are wholly adequate for the practical assessment of a colony as far as physical characters are concerned. Nonetheless, as we have seen, Selection is more than the mere examination of physical characters; only in conjunction with the evidence from performance and ancestry does the assessment of physical characters become a valuable tool of breeding selection. It is to be hoped that as many breeders as possible will avail themselves of this procedure in order to achieve from their efforts the most effective results. In very difficult matters, such as the assessment of drone-producing stocks for the Queen-Mating Stations, specially trained Selection Experts are available to help with essential deci- sions. Conscientiousness, expert knowledge and perseverance are the foundations on which bee breeding must rest.

145 NOTES Intro. 1. Blender. This is the word used in German animal breeding for a hybrid, but it also has overtones of "blinden", to dazzle. So a "blender" is a "shiner", a flash in the pan. 1.1. Heather bee. A form of the Dark Bee, Apis. m.m. lehzeni, adapted to the conditions of the Luneburg Heath, where there was no nectar flow before the heather. Now very mongrelised by Italian imports, very swarmy, and sold to unsuspecting beginners. I 2. Jenter Method while we agree that grafting is simple enough for the ordinary beekeeper, people with defective eyesight have found this method convenient. I 3. The Swiss Grafting Tool mentioned here is obtainable from Steele and Brodie. I 4. Honey Chamber. We have used this term instead of Super, because it applies to a different style of beekeeping, with a second brood box from which the queen may later be excluded and the honey extracted. I 5. Storey Hive. The familiar type with the compartments arranged one above another. It has become popular in Germany only recently, and has not completely displaced other types, the Long Idea, the Leaf Hive, the Trug Hive etc. I 6. Marburg Separating Box. This should be used, as the text implies, when the bees are flying strongly in bright sun. Then the young bees will hurry through the excluder into the dark, while the old bees will fly back to the hive. If used in cool or very dull conditions the bees take much longer to disperse, and a much higher proportion of old bees will enter the box. When the hopper has been closed the box should be put in a cool place and the bees fed. I 7. When the bees are required, the box is jarred or jolted sharply, to dislodge the cluster of bees. The lid is then removed and the bees are immediately sprayed, using an ordinary garden spray (used solely for this purpose) with tepid water. The water will prevent flight, so that the bees can be easily scooped up. If the water is cold the bees will take longer to recover. The quantity of bees required for the BIBBA Mininuc is about 1/3 of a pint. The "cool" place, both for the Marburg box and later for the nuclei, should be about 60° F (15° C). If it is warmer the bees will be restless, if colder the bees can form a "winter" cluster instead of a warm cluster for wax secretion. I 8. Butler Cage. The principle of this cage is to confine the queen without food so that she will readily accept food from the workers. Illustrated and described in "The Encyclopedia of Beekeeping". It is made of woven wire mesh, 6 x 6 to the inch, 22swg wire. This is bent into a tube, 3.1/2 inches long. 3/4 x 1/2 inch in cross section, with one end plugged with wood. The open end is, after the queen has been put in, covered with one thickness of newspaper, held in place with a rubber ring. The cage is placed between two combs with brood. The queen is usually released within 24 hours. Hair curlers can be used in the same way, but whenever hair curlers are used, care should be taken to ensure that the holes are small enough to prevent the entry of workers. We have lost several queens, killed by workers which entered a hair-curler cage. I. 9. Introducing a queen of different race. If she is alone without a nucleus: remove a comb with emerging brood and no bees from the hive, and place it, with the queen, in an empty brood box separated by a screen from the box below. The emerging bees, never having known another queen, will accept her. This process can be continued till the queen has an ample guard, and the two boxes can be united by the paper method. III. 1. Colour markings on mating nucs. See von Frisch: Bees: their vision, chemical senses and language (Cornell University Press) for the shapes and colours most obvious to a bee.

146 III 2. Absconding from nuclei. Insulation is needed to keep bees cool in hot weather (as well as warm in cold weather). Neglect can lead to overheating during a heat wave, when the bees will abscond, even when the queen is mated and has reared brood. Insulate the minicosies with 1 inch expanded polystyrene sheet round the sides, 1.1/2 inches under the lid. An outer protective layer of plywood to prevent damage by birds. Another cause of absconding is unsuitable food, especially hard candy. Follow carefully the recipes given in this book. IV. 1. "Normal" frames. The middle of five sizes of frames in common use in Germany, 370mm x 223mm. IV 2. Colour of the British Bee. It used to be called the "Black Bee", but W. Herrod- Hempsall (Beekeeping New and Old, 1930-1937) denied that a British Black Bee ever existed, and many people today claim that it is brown. We have recently examined several thousand bees, mainly from Yorkshire, and of many which conform to the racial standard for the Mellifera bee we can say: 1. Some appear light brown to the naked eye. This is the colour of the hairs, and close examination reveals black chitin under the hairs. 2. Some appear black to the naked eye. but close examination shows that the second tergite is very dark brown. 3. Some have brown spots or a brown ring on the second tergite but are otherwise black. This is all in accordance with the well known variability of the species. These observations refer to worker bees. A sample of 100 drones from one colony (1987) showed remarkable uniformity in hair colour, Cu I and body colour, all dark, no spots. Only the Saddle Stripe varied, from scarcely visible under magnification to clearly apparent to the naked eye. V. 1. Killing the sample. Killing in a refrigerator is very slow, and leaves the proboscis retracted. Dropping into almost boiling water seems cruel, but is almost instantaneous. The bees should then be dried. When a sample is taken it should be killed as soon as possible. Live bees left to crawl over each other in a small container become coated with nectar etc., and are not fit for examination. V 2. Watchmaker's forceps. Good quality obtainable from H. S. Walsh & Sons Limited 234, Beckenham Road, Beckenham, Kent. BR3 4TS. V 3. Frame wire in the U.K. varies considerably in calibre. Soro Products, Enfield, supplies wire of specified thickness, but only in large quantities. V 4 Mounting wings. Too much sugar will cloud the image of the wing. We advise beginners to take the wing in the forceps by the tip and only dip the other end in the liquid. This will be found adequate to secure the wing in place. V 5 Obtainable in German or English from IBRA V 6 The chart, is available from BIBBA.

147 Artificial Food for Bees

Recipes repeated here for convenience.

Unsuitable food is a common cause of absconding from Mininucs. 1. Schultz Candy. 3 parts finely powdered sugar to one part fluid honey (heated to 60° C against Nosema). 2. Invert Candy. 1kg powdered sugar, 80ml water, 2ml Invertin. Invertin is an enzyme which changes cane sugar into fructose and glucose, so that the candy remains soft. We include this recipe in case Invertin becomes readily available). An invert sugar can readily be made: 1 litre water, plus 2kg sugar plus 4g lactic acid simmered gently for half an hour. (Take care! It froths easily.) lg lactic acid = 0.83ml which can easily be measured with a pipette. This syrup can be used in place of honey in Recipe 1. Other acids which will invert sugar are harmful to bees.

Lactic acid costs about £7 per 500 ml, and with the cost of a pipette for measuring, is much cheaper than honey, carries no disease, and, producing a standard syrup, is much easier to use. lkg syrup = about 0.75 litre, lkg sugar to 0.25 litre syrup is convenient to mix by hand. Powdered sugar = icing sugar without additives. With a little trouble it can be obtained in 50kg bags by a friendly confectioner.

Against Nosema Add lg Fumidil to each Kg candy, mix Fumidil with the dry sugar. Do not add water to the recipe. As Fumidil is highly deliquescent and difficult to weigh in small quantities it is convenient to weigh out 25 oz sugar into a plastic bag, add the contents of a small bottle of Fumidil, mix well, seal and store in a deep freeze 18 degrees F. if possible. When making candy add 1 oz of this mixture for each lg. Fumidil required. If this is done an equal weight should be deducted from the sugar.

148 INDEX

* = illustration

Body markings 113*, 138 Absconding from nuc 87 Discoidal cell 117* Breeder drones 41 Actual Index Value 124f Discoidal shift 35 Breeder, late 139 Age of larvae 17,18* Distribution Breeding 45 ALFONSUS 105 ("scatter") 120, Card 53, 54* Apimondia 12, 53, 75 131* Combination 71 Apis mellifera DREHER 123 Cross 73, 100 carnica 98* Drone-brothers 60 Displacement 76 caucasica 109 Drones Early 139 ligustica 75, 108, 138 body markings 135* for colour 105 mellifera 96, 104*, 138f breeding 41 for production 50 sicula 109 comb 42, 90 frame with trough Arrangement of brood nest 55 Cubital Index 137 feeder 27* Artificial Insemination 71 excluder 89* -line 47, 69 Commercial 41, 63 feeding of 90 material, distribution of 32 instrumental 83* hair colour 136 mood 13 Assessment, quick 118 numbers 88 programme 63-76 Assessment of colony nursing stocks 90. 92 Regulations of German condition 53 physical characters 135 B.K.A. of production 53, 57 -producing colonies 42, 61 Strain 72,104 Average yield 50f DZIERZON 108 systems 22, 29 time 91 time-table 44 Early breeder 139 Basal Vein (of 3rd Cubital value for 8 Economic value 8 cell) 117*, 123* Brood Egg-laying ability 11 BAUMGARTEN 101 Box 28 Egg Queens 20 Bee Colony 59f diseases 99 Emergency mood 21 Bees food 15*, 19*, 21 Exo-skeleton Anatolian 118 food gland 21 colouring 113* Braunelle 107 loss of 65-68 markings 112* Buckfast 98 nest, arrangement of 55 Extra tomentum 116* Carinthian 96 pepperpot 65 Eyepiece with measuring Carniolan 96. 104*, 138* quality of 55, 66* scale 121 Caucasian 109 Brooder colony 23, 26 Country 9, 46 Building urge 58 Dark 96, 104*, 138 FALKENBERG 141f Golden 113 Familiarisation (of plastic Italian 9, 97.103, 138 q-cups 18 Italian hybrids 103 Cage, see Queen-cage Family tree 60 Local 9, 48 Candy, milk 90, 92 Farmers' hives 107 Lower Austrian 105 Carnica see Apis Food reserves 55 Lucerne 145 Chromosome 64 Food, for larvae 15*, 19*, 21 Macedonian 107 Class values (of C.I.) 122-6 Food glands 22 Nigra 96, 108, 139 Colour scale (Goetze) 136 Foot joint 114* Norwegian 100 Colour classes 113 * Foraging zeal 48 Palestinian 100 Colour markings 105,112, Forest-nectar flow 99 Sicilian 109 113* Forewing 117* Slovenian 107 Combination breeding 70f, Swiss 108 78 Bleeding (of stocks) 36 Cordovan Test 82, 93 Gelatine 144 Blender 8, 47, 74* Corner Colony 59 Genetic material, Blood relationship 60,61* Cubital cells 117* combination of 48 BOTTCHER 89,99 Cubital index (C.I.) Glossa 143* Body colour 9, 48, 136 see Index Glossometer 142*

149 GOETZE 99, 136 KOBEL 108 One comb nucleus box Grafting 15, 16f (EWK) 34,39.86 Double- 18 OROSI-PAL 18 Re- 14 Ladle (scoop) 37* Overhair 114*f -tool (Swiss) 16,17* Laidlaw 68 Graticule 112 Larvae grafting of 16, 20 PAGE 68 neglected 14 Pedigree certificate 8 Hair length of 114*, 138 transport of 32 diagram 62* Hair colour 48 Leatherbrown spots 105 Performance Heritable endowment 63 Line breeding 63, 70*, 77* of the colony 53 Heritable value 59 Pure line-breeding 70f of the family 59 Herd-book number 60, 62 Line combination 33 of a single stock 59 HEROLD 115, 126 evaluation of 53 Histogram 128 PESCHETZ 107 Hive Card 53,54* MACKENSEN 64, 145 Petiole 113,135* Hive mother 59 Mandril (for Q-cups) 17 Physical characters Honey production 47, 57 Marburg separating box 36 assessment 8, 42, 52 Humidity 16 Mating examination 111 Humidity chamber 112 Apiary 79, 84 PIANA 29 Hiingler 48 brother-sister 65 Pilosity 114 Husband colonies 60 -flight 83 of tomenta 116*, 139 Hygrometer 16, 31 Nuclei 86f Positive variants 45 Hybrid 51, 73, lOOf Nucleus box 34*. 86* Proboscis length 141 Hybrid stocks 42 setting up 85 actual 144* Hybrid drones 61 Mating Stations 63, 81 measurement 141* Hybrids 51, 73,106 establishment of 85 Production Hybridisation Inland 91, 94 breeding for 50 signs of 105 Island 68, 82 comparison of 48 sneaky 93, 103 Line 81 Productivity Race 84 decline of 46 Matings, further 88 of a colony 53 of a family 53 Inbreeding 47, 63, 65*, 67 Mating weather 92 Progeny, testing of 59 Incompatibility of races 100 MAUL 68, 136 Projector use of 112 Incubator 30 Measuring scale 122* Propolis 99, 109 Index Microscope 112, 120 Provision of food 56 Cubital (C .1.) 48, 117*, Milk candy 90, 92 Pure bred queen, 138 Moritz 68 superiority of 49,51* Mean of C.I. 120,122 Mother-daughter mating 63 Pure breeding 51f C.I. of drone 129 Multi-comb nucleus box 34 area 80, 94 Classes 122f cordon 88, 93. 94* Distribution curve 127f Fan measurement of 118f Negative variant 45* Hantel Index 135 NIEDERWIESER 107 Pre-cubital 135 Nucleus box Quality of brood 65, 66* Radial 135 filling of 37* 86* Queen Instruments 112 Nucleus breeding 12 Introducing cages 31f* "collected" 23 direction finding 85 procedures 38* confinement in cellar 32 excluder 27*, 28* Islands (Spots) 113, 136 Nurse bees 12 for honey production 76 Israel 100 Nursing brood box 28 introduction 38, 41 Nursing colony 12,22* raised from egg 18f preparation of 25*. 27 rearing 12f, 45f KAUSCHE NYE 145 rearing in large batches 29 Killing the test sample 111 working 75f Kirchhainer Mating Box Queen cages 34, 35*. 86 Officially recognised race 108 hair-curler 31*

150 Miller 39, 40* brown 105 One hand 39 yellow 106,108 press on 40* Robbing 109 Test crossings 67 Queen cell 15*, 21*, 31* Royal jelly (queen food) 6 Testing apiary 53 caging 29 runner 99 Thread counter 112,119 cell, built over 30* TIESLER 88 cell utilisation 29 Tomenta 116* cell "wild" 14 Saddle stripes 135 "extra" 116 Queen cup 16,17 Sample 111 pilosity of 138 mandril for 17 Scutellum 113 Tongue see Proboscis Queen mating station Selection 10,18 Tongue reach 139,142* (QMS) 85 Selection for breeding Transport of eggs 37 maintenance of 92 main 8, 111 of larvae and pupae 32 Quickness of development preselection of queen cells 33 49 Selection record 130 TROISECK 73, 106 Quietness on the comb 48, Sex allele 64 50, 58, 139 Sexual maturity 41 Shaken bees see Bleeding Variant, negative 45 SKLENAR 29, 103, 105 Variant, positive 45 Race 7f Slovenia 102 Verification of pure race breeding 76, 94 Special measuring production 53 Races of bees see Apis graticule 121* incompatibility of 94 Spots 113* Watchmakers' forceps 112, Racial hybrids 73 Spring build up 48, 58 144 Racial physical Stakes (for nucs) 86* Wax mirrors 139 characters 112* Starter WEISS 17, 18, 20, 32, 73 standards 137f Colony 23 Wild (natural) queen cells 14 Radial cell 117 Foundation 91 WOHLGEMUTH 39, 49, 50 Rapidity of build up 48 Stimulative feeding 43, 90 Working queens 75f Readiness for breeding 12 Strains WOYKE 64 Rearing of drones 41 Braunelle 107 WRISNIG 106 Queens final stage 23 Bukovsek 73 from the egg 18 Peschetz 106 in de-queened stock 23 Sklenar 73,106 Yield in Queen-right stock 24f, Troiseck 73,106 nett 47*, 50, 57 27* STRGAR 107 percentage 57 Red clover Swarming 56, 57 Re-queening, rapid 39 inclination to 48, 139 Reserve stocks 35, 91 intention to 12 ZANDER 17, 49, 108 Resourcefulness 48 mood (urge) 29, 58 Rings 113* prevention 36

151